From 2e34ab007fd9ad99c11164325749c360626e01d3 Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 15 Jul 2013 10:54:24 +0100 Subject: [PATCH 01/18] Edited B122 --- 10.3897_zookeys.316.5132.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index 699069d..af671fe 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -705,7 +705,7 @@ Villeneuve J (1933) Contribution a la classification des Tachinariae paléarctiques. Ve Congrès International d’Entomologie, Paris, 18–24 juillet 1932, 2: 243–255. Wainwright CJ (1928) The British Tachinidae (Diptera). Transactions of the Entomological Society of London 76: 139–254 + pls. IX–X. Wood DM (1972) A revision of the New World Exoristini (Diptera: Tachinidae). I. Phorocera subgenus Pseudotachinomyia. Canadian Entomologist 104: 471-503. - Wood DM (1985) A taxonomic conspectus of the Blondeliini of North and Central America and the West Indies (Diptera: Tachinidae). Memoirs of the Entomological Society of Canada 132: 130 pp. + Wood DM (1985) A taxonomic conspectus of the Blondeliini of North and Central America and the West Indies (Diptera: Tachinidae). Memoirs of the Entomological Society of Canada 132: 130 pp. doi: 10.4039/entm117132fv Wood DM (1987) Tachinidae. In: McAlpine JFPeterson BVShewell GETeskey HJVockeroth JRWood DM (Eds). Manual of Nearctic Diptera. Volume 2. Agriculture Canada Monograph 28: 1193–1269. Wood DMZumbado MA (2010) Tachinidae (tachinid flies, parasitic flies). In: Brown BVBorkent ACumming JMWood DMWoodley NEZumbado MA (Eds). Manual of Central American Diptera. Volume 2. NRC Research Press, Ottawa: 1343-1417. From 251dafbd96d1e81f4bde256a4582cfa7cd6b95fa Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 15 Jul 2013 10:57:37 +0100 Subject: [PATCH 02/18] Deleted extra --- 10.3897_zookeys.316.5132.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index af671fe..5294b12 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -705,7 +705,7 @@ Villeneuve J (1933) Contribution a la classification des Tachinariae paléarctiques. Ve Congrès International d’Entomologie, Paris, 18–24 juillet 1932, 2: 243–255. Wainwright CJ (1928) The British Tachinidae (Diptera). Transactions of the Entomological Society of London 76: 139–254 + pls. IX–X. Wood DM (1972) A revision of the New World Exoristini (Diptera: Tachinidae). I. Phorocera subgenus Pseudotachinomyia. Canadian Entomologist 104: 471-503. - Wood DM (1985) A taxonomic conspectus of the Blondeliini of North and Central America and the West Indies (Diptera: Tachinidae). Memoirs of the Entomological Society of Canada 132: 130 pp. doi: 10.4039/entm117132fv + Wood DM (1985) A taxonomic conspectus of the Blondeliini of North and Central America and the West Indies (Diptera: Tachinidae). Memoirs of the Entomological Society of Canada 132: 130 pp. doi: 10.4039/entm117132fv Wood DM (1987) Tachinidae. In: McAlpine JFPeterson BVShewell GETeskey HJVockeroth JRWood DM (Eds). Manual of Nearctic Diptera. Volume 2. Agriculture Canada Monograph 28: 1193–1269. Wood DMZumbado MA (2010) Tachinidae (tachinid flies, parasitic flies). In: Brown BVBorkent ACumming JMWood DMWoodley NEZumbado MA (Eds). Manual of Central American Diptera. Volume 2. NRC Research Press, Ottawa: 1343-1417. From e061c6607722573e8abd529543d5052f684f37b3 Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 15 Jul 2013 11:02:11 +0100 Subject: [PATCH 03/18] Parsed B92 and added handle --- 10.3897_zookeys.316.5132.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index 5294b12..1f471b9 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -673,7 +673,7 @@ Sabrosky CW (1961) Rondani’s “Dipterologiae italicae prodromus”. Annals of the Entomological Society of America 54: 827-831. Sabrosky CW (1999) Family-group names in Diptera. An annotated catalog. Myia 10: 360 pp. Sabrosky CWArnaud PH Jr (1965) Family Tachinidae (Larvaevoridae). Pp. 961–1108. In: Stone A, Sabrosky CW, Wirth WW, Foote RH, Coulson JR (Eds) A catalog of the Diptera of America north of Mexico. United States Department of Agriculture. Agriculture Handbook 276: iv + 1696 pp. - Sasaki C (1887) On the life-history of Ugimya sericaria, Rondani. Journal of the College of Science, Imperial University, Japan 1: 1–46 + 6 pls. + Sasaki C (1887) On the life-history of Ugimya sericaria, Rondani. Journal of the College of Science, Imperial University, Japan 1: 1–46 + 6 pls. handle: 2261/28848 Schiner JR (1860–1864) Fauna Austriaca. Die Fliegen (Diptera). Nach der analytischen Methode bearbeitet, mit der Charakteristik sämmtlicher europäischer Gattungen, der Beschreibung aller in Deutschland vorkommenden Arten und der Aufzählung aller bisher beschriebenen europäischen Arten. C Gerold’s Sohn, Wien. lxxx + 2 pls. + 674 pp. Shima H (1983) A new species of Oxyphyllomyia (Diptera, Tachinidae) from Nepal, with reference to the phylogenetic position of the genus. Annotationes Zoologicae Japonenses 56: 338-350. From f9f04633618ead262c24210b460e4852782405db Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 15 Jul 2013 11:03:02 +0100 Subject: [PATCH 04/18] B92 --- 10.3897_zookeys.316.5132.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index 1f471b9..2c260b1 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -673,7 +673,7 @@ Sabrosky CW (1961) Rondani’s “Dipterologiae italicae prodromus”. Annals of the Entomological Society of America 54: 827-831. Sabrosky CW (1999) Family-group names in Diptera. An annotated catalog. Myia 10: 360 pp. Sabrosky CWArnaud PH Jr (1965) Family Tachinidae (Larvaevoridae). Pp. 961–1108. In: Stone A, Sabrosky CW, Wirth WW, Foote RH, Coulson JR (Eds) A catalog of the Diptera of America north of Mexico. United States Department of Agriculture. Agriculture Handbook 276: iv + 1696 pp. - Sasaki C (1887) On the life-history of Ugimya sericaria, Rondani. Journal of the College of Science, Imperial University, Japan 1: 1–46 + 6 pls. handle: 2261/28848 + Sasaki C (1887) On the life-history of Ugimya sericaria, Rondani. Journal of the College of Science, Imperial University, Japan 1: 1–46 + 6 pls. handle: 2261/28848 Schiner JR (1860–1864) Fauna Austriaca. Die Fliegen (Diptera). Nach der analytischen Methode bearbeitet, mit der Charakteristik sämmtlicher europäischer Gattungen, der Beschreibung aller in Deutschland vorkommenden Arten und der Aufzählung aller bisher beschriebenen europäischen Arten. C Gerold’s Sohn, Wien. lxxx + 2 pls. + 674 pp. Shima H (1983) A new species of Oxyphyllomyia (Diptera, Tachinidae) from Nepal, with reference to the phylogenetic position of the genus. Annotationes Zoologicae Japonenses 56: 338-350. From 8a801c594423dd5f25b2987f0cd2077b2cf29d32 Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 15 Jul 2013 11:03:50 +0100 Subject: [PATCH 05/18] B92 --- 10.3897_zookeys.316.5132.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index 2c260b1..843bbc1 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -673,7 +673,7 @@ Sabrosky CW (1961) Rondani’s “Dipterologiae italicae prodromus”. Annals of the Entomological Society of America 54: 827-831. Sabrosky CW (1999) Family-group names in Diptera. An annotated catalog. Myia 10: 360 pp. Sabrosky CWArnaud PH Jr (1965) Family Tachinidae (Larvaevoridae). Pp. 961–1108. In: Stone A, Sabrosky CW, Wirth WW, Foote RH, Coulson JR (Eds) A catalog of the Diptera of America north of Mexico. United States Department of Agriculture. Agriculture Handbook 276: iv + 1696 pp. - Sasaki C (1887) On the life-history of Ugimya sericaria, Rondani. Journal of the College of Science, Imperial University, Japan 1: 1–46 + 6 pls. handle: 2261/28848 + Sasaki C (1887) On the life-history of Ugimya sericaria, Rondani. Journal of the College of Science, Imperial University, Japan 1: 1–46 + 6 pls. handle: 2261/28848 Schiner JR (1860–1864) Fauna Austriaca. Die Fliegen (Diptera). Nach der analytischen Methode bearbeitet, mit der Charakteristik sämmtlicher europäischer Gattungen, der Beschreibung aller in Deutschland vorkommenden Arten und der Aufzählung aller bisher beschriebenen europäischen Arten. C Gerold’s Sohn, Wien. lxxx + 2 pls. + 674 pp. Shima H (1983) A new species of Oxyphyllomyia (Diptera, Tachinidae) from Nepal, with reference to the phylogenetic position of the genus. Annotationes Zoologicae Japonenses 56: 338-350. From 9018d1525979f2d5e6451787383c4e657a99e4df Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 15 Jul 2013 20:00:36 +0100 Subject: [PATCH 06/18] B8 --- 10.3897_zookeys.316.5132.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index 843bbc1..d352acd 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -584,7 +584,7 @@ Belshaw R (1993) Tachinid flies. Diptera: Tachinidae. Handbooks for the identification of British Insects, 10, Part 4a(i). Royal Entomological Society of London, London. 169 pp. Bezzi MStein P (1907) Cyclorrapha Aschiza. Cyclorrapha Schizophora: Schizometopa. Pp. 1–747. In: Becker T, Bezzi M, Kertész K, Stein P (Eds) Katalog der paläarktischen Dipteren. Band III. Budapest. 828 pp. Brauer FBergenstamm JE von (1889–1895) Die Zweiflügler des Kaiserlichen Museums zu Wien. IV–VII. Vorarbeiten zu einer Monographie der Muscaria Schizometopa (exclusive Anthomyidae). Partes I– IV. F Tempsky, Wien. - Cantrell BK (1984) Synopsis of the Australian Phasiinae, including revisions of Gerocyptera Townsend and the Australian species of Cylindromyia Meigen (Diptera: Tachinidae). Australian Journal of Zoology. Supplementary Series 102: 1-60. + Cantrell BK (1984) Synopsis of the Australian Phasiinae, including revisions of Gerocyptera Townsend and the Australian species of Cylindromyia Meigen (Diptera: Tachinidae). Australian Journal of Zoology. Supplementary Series 102: 1-60. doi: 10.1071/AJZS102 Cantrell BK (1988) The comparative morphology of the male and female postabdomen of the Australian Tachinidae (Diptera), with descriptions of some first-instar larvae and pupae. Invertebrate Taxonomy 2: 81-221. Cantrell BKBurwell CJ (2010) The tribe Dufouriini (Diptera: Tachinidae: Dexiinae) recorded from Australia with the description of two new species. Memoirs of the Queensland Museum 55: 119-133. From 5f7d66c71bc75b8456ef557d0d0fc4613b18a26f Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Wed, 17 Jul 2013 10:26:29 +0100 Subject: [PATCH 07/18] DOI added for B4 using script --- 10.3897_zookeys.316.5132.xml | 448 +++++++++++++++++------------------ 1 file changed, 224 insertions(+), 224 deletions(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index d352acd..51e4820 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -1,6 +1,6 @@  -
+
ZooKeys @@ -22,7 +22,7 @@ - O’Hara + O’Hara James E. 1 @@ -33,7 +33,7 @@ -

Corresponding author: James E. O’Hara (james.ohara@agr.gc.ca)

+

Corresponding author: James E. O’Hara (james.ohara@agr.gc.ca)

Academic editor: P. Cerretti

@@ -63,7 +63,7 @@ - James E. O’Hara + James E. O’Hara This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. @@ -86,62 +86,62 @@ Introduction

The Tachinidae are among the largest families of Diptera with about 8500 valid species1. One can only guess at the true diversity of the family but at least double the number of valid species is a conservative estimate. What is not in doubt is the important ecological role these parasitoid flies play in the environment. It is desirable to organize these flies into a phylogenetically stable suprageneric classification as an aid to those who study them and to enable predictions to be made about the less studied species based on the known habits of related species.

-

The Tachinidae may not be the single largest family of flies on Earth but in terms of genera they tower over all of the other 140-odd families. The current number of valid genera is about 1520 (O’Hara 2012). The next largest family is Cecidomyiidae with about 760 genera and there are only two other families with more than 500 genera: Asilidae and Chironomidae (Pape et al. 2011). Taxonomically the Tachinidae are arguably the most difficult family of flies and perhaps because of this plus the size of the family and their high position on the evolutionary tree of Diptera they have received scant attention below the family level by those investigating dipteran relationships (e.g., Yeates et al. 2007, Kutty et al. 2010). There is currently an international effort aimed at addressing this imbalance by specifically targeting the Tachinidae for phylogenetic analysis using morphological and molecular data (Stireman et al. 2013).

+

The Tachinidae may not be the single largest family of flies on Earth but in terms of genera they tower over all of the other 140-odd families. The current number of valid genera is about 1520 (O’Hara 2012). The next largest family is Cecidomyiidae with about 760 genera and there are only two other families with more than 500 genera: Asilidae and Chironomidae (Pape et al. 2011). Taxonomically the Tachinidae are arguably the most difficult family of flies and perhaps because of this plus the size of the family and their high position on the evolutionary tree of Diptera they have received scant attention below the family level by those investigating dipteran relationships (e.g., Yeates et al. 2007, Kutty et al. 2010). There is currently an international effort aimed at addressing this imbalance by specifically targeting the Tachinidae for phylogenetic analysis using morphological and molecular data (Stireman et al. 2013).

It seems appropriate at this time to review the history of tachinid classification from its earliest beginnings, tracing how it has changed in response to discoveries of phylogenetically insightful characters and was affected by conflicting views on the nature of generic and suprageneric limits. The noticeable disharmony in the way tachinids were classified among the six biogeographic regions of the world is still in evidence today. The task that now awaits present and future tachinidologists is to determine to a better degree than in the past the evolutionary history of the Tachinidae and to classify the family in a manner than reflects its phylogeny and preserves the best elements of the most recent classifications.

 The early years -

The meagre number of tachinid species known in the early 1800s was placed in about a dozen genera with the majority of them in Meigen’s (1803) broadly defined Tachina2. André-Jean-Baptiste Robineau-Desvoidy revolutionized tachinid classification with the publication of his Essai sur les Myodaires (Robineau-Desvoidy 1830), in which approximately 130 new genera now placed in Tachinidae were described (Evenhuis et al. 2010). Of this total, 73 genera are presently treated as valid (O’Hara 2012). Robineau-Desvoidy (1830) also proposed the name “Calypteratae” (Calyptratae) for a higher group within his Myodaria (essentially modern-day Schizophora), which with some modification in concept (most notably the addition of the Anthomyiidae) is now regarded as “one of the best established monophyletic subsections of the Schizophora” (McAlpine 1989: 1425). In this same work, Robineau-Desvoidy’s six tribes of Calypteratae approximated some of the familial and subfamilial groupings in use today in this subsection. One of these, the Entomobiae (including most of the then-known taxa of the Tachinidae), included a small number of genera grouped under the Tachinariae. The priority of the name Tachinidae over other family-group names available for this family thus dates from Robineau-Desvoidy (1830).

-

The Essai sur les Myodaires was not without its faults and received mixed reviews from dipterists of the day. Robineau-Desvoidy’s final contribution to dipterology, a massive two-volume work published in 1863 and six years after his death, Histoire naturelle des diptères des environs de Paris, has been justly criticized as an inferior work. In it were proposed about 160 new tachinid genera, only 25 of which are currently recognized as valid (O’Hara 2012). Similarly, a huge number of new species were described with many of them later becoming junior synonyms or nomina dubia (the latter resulting from the destruction of many of Robineau-Desvoidy’s name-bearing types, Evenhuis et al. 2010: 233).

-

Contemporaneous with Robineau-Desvoidy were Meigen, Wiedemann, Macquart and Walker, each of whom contributed significantly to the description of species but not much to the higher classification of what are now the Tachinidae. Macquart (e.g., Diptères exotiques nouveaux ou peu connus, 1838–1855), like Robineau-Desvoidy, recognized the need for more genera to accommodate the emerging diversity and throughout his career described about 100 tachinid genera, of which 45 are currently valid (O’Hara 2012).

-

The study of Diptera during the mid to late 1800s continued to be led by Europeans. Among the more notable achievements during this time were the regional treatments on the Diptera of Scandinavia by Zetterstedt (1842–1860), on Italian Diptera by Rondani (1856–1880), and on Austrian Diptera by Schiner (1860–1864). A most ambitious and influential work on the Diptera of the Vienna Museum by Brauer and Bergenstamm (1889–1895) contributed greatly to the knowledge of world Tachinidae, but was marred by an unsatisfactory and artificial suprageneric classification (e.g., Coquillett 1897, Aldrich 1905, Villeneuve 1924, Wainwright 1928, Mesnil 1944). Brauer and Bergenstamm described over 250 genera and subgenera of Tachinidae, of which 99 are currently valid genera (O’Hara 2012).

-

New World tachinids came under increased attention near the turn of the century, first by van der Wulp (1888–1891) and then by Coquillett (1897). Commenting on the state of tachinid classification at the time, Coquillett (1897: 27) noted:

-

“Probably no single family of Diptera has received greater consideration in Europe than the Tachinidae, and yet, strange as this may seem, no other family at the present time is in greater disorder. Several authors accord them only subfamily rank, but it appears desirable to consider them as a distinct family, although their relationship to the Dexidae and Sarcophagidae is a very intimate one.”

-

Coquillett (1897) recognized five subfamilies of Tachinidae, four representing present-day Phasiinae and one (his Tachininae) representing modern Exoristinae + Tachininae. No tribes were recognized. The “Dexidae” (Dexiidae) were regarded as a separate family and excluded.

-

Despite the chaotic state of tachinid classification in the late 1800s, an important methodological advance was made in the study of dipteran characters that would lead to a better understanding of natural groupings within the higher Diptera. Early authors like Meigen, Macquart and Robineau-Desvoidy had used certain large setae in their descriptions but it was Rondani (1845) who would apply the term macrochaetae (as “macrochetae”) to them. Later Osten Sacken (1881, 1884) would formalize a nomenclature for such macrochaetae under the term chaetotaxy. With refinements of the system by Girschner (1893, 1896), the study of chaetotaxy began to revolutionize the study of the more setose Diptera. Osten Sacken (1884: 511) had observed that the “hypopleural” (meral) setae “occur only in some of the Diptera Calyptrata, which have a row or a tuft of them” and Girschner (1893) used this characteristic to define the Tachinidae in the broad sense of present-day Oestroidea. Girschner also recognized several subgroups within Tachinidae s. lat. based on other setal arrangements. The classification was not completely satisfactory and it was only later that the enlarged subscutellum would be used to delimit the Tachinidae in a more modern sense (see below). Frey (1921) built upon the work of Girschner to further advance the classification of this group of flies.

-

By the beginning of the 20th Century the taxonomic literature on Palaearctic Diptera was both voluminous and daunting, especially for new students of the group. The Katalog der paläarktischen Dipteren (1903–1907) was therefore of immense importance, bringing together under a single classification all the names of Palaearctic Diptera. The part by Bezzi and Stein (1907) on the Schizometopa relied heavily on the work of Girschner and proposed a higher classification of considerable merit for its day. The Schizometopa were split into two families, Tachinidae and “Anthomyidae” (Anthomyiidae) (present-day Muscoidea). Within Tachinidae, ten subfamilies were recognized and listed in the following order: Tachininae, Dexiinae, Rhinophorinae, Sarcophaginae, Calliphorinae, Phasiinae, Eginiinae, “Hypoderminae” (Hypodermatinae), Oestrinae and “Gastrophilinae” (Gasterophilinae). With the exception of the Eginiinae (now placed in Muscidae), the rest of the groups with some adjustment to relative ranking closely approximates the families now recognized in the Oestroidea.

-

Although the Katalog der paläarktischen Dipteren must have been a most welcome addition to the shelf of any dipterist of the day, Mesnil (1944: 2) later criticized the Bezzi and Stein (1907) portion on the grounds that it was “voll von Irrtümern und praktisch unverwendbar” [“full of mistakes and practically unusable”].

+

The meagre number of tachinid species known in the early 1800s was placed in about a dozen genera with the majority of them in Meigen’s (1803) broadly defined Tachina2. André-Jean-Baptiste Robineau-Desvoidy revolutionized tachinid classification with the publication of his Essai sur les Myodaires (Robineau-Desvoidy 1830), in which approximately 130 new genera now placed in Tachinidae were described (Evenhuis et al. 2010). Of this total, 73 genera are presently treated as valid (O’Hara 2012). Robineau-Desvoidy (1830) also proposed the name “Calypteratae” (Calyptratae) for a higher group within his Myodaria (essentially modern-day Schizophora), which with some modification in concept (most notably the addition of the Anthomyiidae) is now regarded as “one of the best established monophyletic subsections of the Schizophora” (McAlpine 1989: 1425). In this same work, Robineau-Desvoidy’s six tribes of Calypteratae approximated some of the familial and subfamilial groupings in use today in this subsection. One of these, the Entomobiae (including most of the then-known taxa of the Tachinidae), included a small number of genera grouped under the Tachinariae. The priority of the name Tachinidae over other family-group names available for this family thus dates from Robineau-Desvoidy (1830).

+

The Essai sur les Myodaires was not without its faults and received mixed reviews from dipterists of the day. Robineau-Desvoidy’s final contribution to dipterology, a massive two-volume work published in 1863 and six years after his death, Histoire naturelle des diptères des environs de Paris, has been justly criticized as an inferior work. In it were proposed about 160 new tachinid genera, only 25 of which are currently recognized as valid (O’Hara 2012). Similarly, a huge number of new species were described with many of them later becoming junior synonyms or nomina dubia (the latter resulting from the destruction of many of Robineau-Desvoidy’s name-bearing types, Evenhuis et al. 2010: 233).

+

Contemporaneous with Robineau-Desvoidy were Meigen, Wiedemann, Macquart and Walker, each of whom contributed significantly to the description of species but not much to the higher classification of what are now the Tachinidae. Macquart (e.g., Diptères exotiques nouveaux ou peu connus, 1838–1855), like Robineau-Desvoidy, recognized the need for more genera to accommodate the emerging diversity and throughout his career described about 100 tachinid genera, of which 45 are currently valid (O’Hara 2012).

+

The study of Diptera during the mid to late 1800s continued to be led by Europeans. Among the more notable achievements during this time were the regional treatments on the Diptera of Scandinavia by Zetterstedt (1842–1860), on Italian Diptera by Rondani (1856–1880), and on Austrian Diptera by Schiner (1860–1864). A most ambitious and influential work on the Diptera of the Vienna Museum by Brauer and Bergenstamm (1889–1895) contributed greatly to the knowledge of world Tachinidae, but was marred by an unsatisfactory and artificial suprageneric classification (e.g., Coquillett 1897, Aldrich 1905, Villeneuve 1924, Wainwright 1928, Mesnil 1944). Brauer and Bergenstamm described over 250 genera and subgenera of Tachinidae, of which 99 are currently valid genera (O’Hara 2012).

+

New World tachinids came under increased attention near the turn of the century, first by van der Wulp (1888–1891) and then by Coquillett (1897). Commenting on the state of tachinid classification at the time, Coquillett (1897: 27) noted:

+

“Probably no single family of Diptera has received greater consideration in Europe than the Tachinidae, and yet, strange as this may seem, no other family at the present time is in greater disorder. Several authors accord them only subfamily rank, but it appears desirable to consider them as a distinct family, although their relationship to the Dexidae and Sarcophagidae is a very intimate one.”

+

Coquillett (1897) recognized five subfamilies of Tachinidae, four representing present-day Phasiinae and one (his Tachininae) representing modern Exoristinae + Tachininae. No tribes were recognized. The “Dexidae” (Dexiidae) were regarded as a separate family and excluded.

+

Despite the chaotic state of tachinid classification in the late 1800s, an important methodological advance was made in the study of dipteran characters that would lead to a better understanding of natural groupings within the higher Diptera. Early authors like Meigen, Macquart and Robineau-Desvoidy had used certain large setae in their descriptions but it was Rondani (1845) who would apply the term macrochaetae (as “macrochetae”) to them. Later Osten Sacken (1881, 1884) would formalize a nomenclature for such macrochaetae under the term chaetotaxy. With refinements of the system by Girschner (1893, 1896), the study of chaetotaxy began to revolutionize the study of the more setose Diptera. Osten Sacken (1884: 511) had observed that the “hypopleural” (meral) setae “occur only in some of the Diptera Calyptrata, which have a row or a tuft of them” and Girschner (1893) used this characteristic to define the Tachinidae in the broad sense of present-day Oestroidea. Girschner also recognized several subgroups within Tachinidae s. lat. based on other setal arrangements. The classification was not completely satisfactory and it was only later that the enlarged subscutellum would be used to delimit the Tachinidae in a more modern sense (see below). Frey (1921) built upon the work of Girschner to further advance the classification of this group of flies.

+

By the beginning of the 20th Century the taxonomic literature on Palaearctic Diptera was both voluminous and daunting, especially for new students of the group. The Katalog der paläarktischen Dipteren (1903–1907) was therefore of immense importance, bringing together under a single classification all the names of Palaearctic Diptera. The part by Bezzi and Stein (1907) on the Schizometopa relied heavily on the work of Girschner and proposed a higher classification of considerable merit for its day. The Schizometopa were split into two families, Tachinidae and “Anthomyidae” (Anthomyiidae) (present-day Muscoidea). Within Tachinidae, ten subfamilies were recognized and listed in the following order: Tachininae, Dexiinae, Rhinophorinae, Sarcophaginae, Calliphorinae, Phasiinae, Eginiinae, “Hypoderminae” (Hypodermatinae), Oestrinae and “Gastrophilinae” (Gasterophilinae). With the exception of the Eginiinae (now placed in Muscidae), the rest of the groups with some adjustment to relative ranking closely approximates the families now recognized in the Oestroidea.

+

Although the Katalog der paläarktischen Dipteren must have been a most welcome addition to the shelf of any dipterist of the day, Mesnil (1944: 2) later criticized the Bezzi and Stein (1907) portion on the grounds that it was “voll von Irrtümern und praktisch unverwendbar” [“full of mistakes and practically unusable”].

Classifying New World <tp:taxon-name>Tachinidae</tp:taxon-name> -

North American Diptera were first catalogued by Osten Sacken (1858) and the few tachinid genera listed therein were included in the Muscidae. In the second edition of his catalogue, Osten Sacken (1878) revised the classification of Diptera and recognized both the Tachinidae and “Dexidae” (Dexiidae) as families. The next catalogue was that of Aldrich (1905), and although the Tachinidae and Dexiidae were kept separate following Osten Sacken (1878) and Coquillett (1897), the suggestion was made that they might be better combined. Aldrich (1905) followed the order of genera of Tachinidae given by Coquillett (1897) and interpolated additional genera and species as necessary. Disparaging remarks were made about the monographic works of van der Wulp (1888–1891) and Brauer and Bergenstamm (1889–1895), and of the species descriptions of Bigot (“in every way objectionable, almost always referred to the wrong genus, and seldom containing the essential data”, Aldrich 1905: 420).

-

Charles Henry Tyler Townsend, the most eccentric and prolific of all tachinidologists, published his first paper on tachinids in 1891 and his last in 1944, with almost 500 publications in total (the majority on tachinids) over this long period (Arnaud 1958). He took up the study of insects at the age of 10 and the study of flies at 25. He held a variety of jobs and professional appointments in the United States and later Peru before settling in Brazil for the last 25 years of his life (Townsend 1943). His most significant achievement was the Manual of Myiology, a 12-volume series on the “Oestromuscaria” published between 1934–1942 in which virtually every genus of these flies known at the time was placed in a suprageneric classification and given a detailed description.

-

Townsend was, by his own admission, a splitter of taxa. He was well versed in the works of others and offered this historical perspective on the struggle between “radicalism and conservatism” (Townsend 1935: 37):

-

“History shows that the taxonomy of these flies has suffered much in the making, subjected as it has been for the past century to an alternation of radicalism and conservatism, commonly called splitting and lumping. … Desvoidy, the first radical, employed restricted genera and Macquart, the first conservative, lumped them; Róndani again restricted the genera and Schiner lumped them; Brauer & Bergenstamm split, while Coquillett and Aldrich lumped; Villeneuve and Townsend again split, while Curran and Malloch lumped.”

-

The restricted genera of Townsend were based on the author’s concept of a “physiological genus”, defined as a “natural genus” comprising “all those species which can produce fertile crosses” (Townsend 1935: 38). As noted by van Emden (1945: 389–390), “the adoption of [this] principle implies the application of the generic unit to every unit considered to be a species in general zoological practice”. One can learn, explained Townsend (1935: 56), “to make a complete description of a fly genus and its genotype [type species] in one hour for one sex and an hour and a half for both sexes”. The ideal number of members within each of the categories of genus, tribe, family, suborder and order was set at five (Townsend 1935: 60–61). In practise Townsend rarely included more than one species per genus and throughout his career described 1491 genera and 1555 species (Arnaud 1958), with approximately 85% of the genera belonging to the Tachinidae. The number of valid tachinid genera attributed to Townsend currently stands at 544 (O’Hara 2012), more than five times that of any other author. Second place is held by Robineau-Desvoidy with 104 valid genera (O’Hara 2012).

-

Townsend’s methods and productivity are worth more than a cursory mention because this author has, in some ways, done more to retard tachinid taxonomy than advance it. The sheer volume of genera is one problem, and their assignment to suprageneric categories is another. Townsend knew that females of Tachinidae and related families possess a great diversity of reproductive systems that produce different kinds of eggs and larvae. After 25 years of dissecting specimens and studying the female reproductive system, he was able to recognize 36 distinct groups, most pertaining to present-day Tachinidae (Townsend 1934). Townsend (1935: 38) believed that tachinid relationships had proved to be a “Gordian knot” in the past and:

-

“not until the wonderful diversity of female reproductive characters and early stages was demonstrated did any sword for the cutting of this knot appear. … We are now able to determine actual relationships with greater certainty, having found the key to affinities by correlating external anatomic characters with internal reproductive and early stage characters.”

-

Thus armed with internal, external and larval characters, Townsend developed a unique classification that divided present-day Tachinidae among seven families (Gymnosomatidae, Oestridae, Prosenidae, Rutiliidae, Tachinidae, Dexiidae, Exoristidae) and about 90 tribes. Had this hierarchical system truly classified the Tachinidae along phylogenetic lines then it would have been the most significant advance in the history of tachinidology. However, it fell short of this goal and is now regarded as both unmanageable and artificial (e.g., Mesnil 1939, Wood 1985, 1987). Specialists also found the keys to tribes and genera in Manual of Myiology to be fraught with problems, thus hindering the recognition of Townsend’s supraspecific taxa.

-

William Robin Thompson published a series of eight papers in the Tachinids of Trinidad (Thompson 1961–1968). He had difficulty interpreting the fauna of Trinidad according to the Townsend scheme and chose to avoid attempting to revise Townsend’s genera:

-

“[I have] decided also that in most cases an attempt to simplify the taxonomic problems by reducing Townsendian genera to the synonymy is impracticable because with the knowledge we now have it is impossible to know when to stop” (Thompson 1961: 22).

+

North American Diptera were first catalogued by Osten Sacken (1858) and the few tachinid genera listed therein were included in the Muscidae. In the second edition of his catalogue, Osten Sacken (1878) revised the classification of Diptera and recognized both the Tachinidae and “Dexidae” (Dexiidae) as families. The next catalogue was that of Aldrich (1905), and although the Tachinidae and Dexiidae were kept separate following Osten Sacken (1878) and Coquillett (1897), the suggestion was made that they might be better combined. Aldrich (1905) followed the order of genera of Tachinidae given by Coquillett (1897) and interpolated additional genera and species as necessary. Disparaging remarks were made about the monographic works of van der Wulp (1888–1891) and Brauer and Bergenstamm (1889–1895), and of the species descriptions of Bigot (“in every way objectionable, almost always referred to the wrong genus, and seldom containing the essential data”, Aldrich 1905: 420).

+

Charles Henry Tyler Townsend, the most eccentric and prolific of all tachinidologists, published his first paper on tachinids in 1891 and his last in 1944, with almost 500 publications in total (the majority on tachinids) over this long period (Arnaud 1958). He took up the study of insects at the age of 10 and the study of flies at 25. He held a variety of jobs and professional appointments in the United States and later Peru before settling in Brazil for the last 25 years of his life (Townsend 1943). His most significant achievement was the Manual of Myiology, a 12-volume series on the “Oestromuscaria” published between 1934–1942 in which virtually every genus of these flies known at the time was placed in a suprageneric classification and given a detailed description.

+

Townsend was, by his own admission, a splitter of taxa. He was well versed in the works of others and offered this historical perspective on the struggle between “radicalism and conservatism” (Townsend 1935: 37):

+

“History shows that the taxonomy of these flies has suffered much in the making, subjected as it has been for the past century to an alternation of radicalism and conservatism, commonly called splitting and lumping. … Desvoidy, the first radical, employed restricted genera and Macquart, the first conservative, lumped them; Róndani again restricted the genera and Schiner lumped them; Brauer & Bergenstamm split, while Coquillett and Aldrich lumped; Villeneuve and Townsend again split, while Curran and Malloch lumped.”

+

The restricted genera of Townsend were based on the author’s concept of a “physiological genus”, defined as a “natural genus” comprising “all those species which can produce fertile crosses” (Townsend 1935: 38). As noted by van Emden (1945: 389–390), “the adoption of [this] principle implies the application of the generic unit to every unit considered to be a species in general zoological practice”. One can learn, explained Townsend (1935: 56), “to make a complete description of a fly genus and its genotype [type species] in one hour for one sex and an hour and a half for both sexes”. The ideal number of members within each of the categories of genus, tribe, family, suborder and order was set at five (Townsend 1935: 60–61). In practise Townsend rarely included more than one species per genus and throughout his career described 1491 genera and 1555 species (Arnaud 1958), with approximately 85% of the genera belonging to the Tachinidae. The number of valid tachinid genera attributed to Townsend currently stands at 544 (O’Hara 2012), more than five times that of any other author. Second place is held by Robineau-Desvoidy with 104 valid genera (O’Hara 2012).

+

Townsend’s methods and productivity are worth more than a cursory mention because this author has, in some ways, done more to retard tachinid taxonomy than advance it. The sheer volume of genera is one problem, and their assignment to suprageneric categories is another. Townsend knew that females of Tachinidae and related families possess a great diversity of reproductive systems that produce different kinds of eggs and larvae. After 25 years of dissecting specimens and studying the female reproductive system, he was able to recognize 36 distinct groups, most pertaining to present-day Tachinidae (Townsend 1934). Townsend (1935: 38) believed that tachinid relationships had proved to be a “Gordian knot” in the past and:

+

“not until the wonderful diversity of female reproductive characters and early stages was demonstrated did any sword for the cutting of this knot appear. … We are now able to determine actual relationships with greater certainty, having found the key to affinities by correlating external anatomic characters with internal reproductive and early stage characters.”

+

Thus armed with internal, external and larval characters, Townsend developed a unique classification that divided present-day Tachinidae among seven families (Gymnosomatidae, Oestridae, Prosenidae, Rutiliidae, Tachinidae, Dexiidae, Exoristidae) and about 90 tribes. Had this hierarchical system truly classified the Tachinidae along phylogenetic lines then it would have been the most significant advance in the history of tachinidology. However, it fell short of this goal and is now regarded as both unmanageable and artificial (e.g., Mesnil 1939, Wood 1985, 1987). Specialists also found the keys to tribes and genera in Manual of Myiology to be fraught with problems, thus hindering the recognition of Townsend’s supraspecific taxa.

+

William Robin Thompson published a series of eight papers in the Tachinids of Trinidad (Thompson 1961–1968). He had difficulty interpreting the fauna of Trinidad according to the Townsend scheme and chose to avoid attempting to revise Townsend’s genera:

+

“[I have] decided also that in most cases an attempt to simplify the taxonomic problems by reducing Townsendian genera to the synonymy is impracticable because with the knowledge we now have it is impossible to know when to stop” (Thompson 1961: 22).

Thompson found the works of Mesnil and other Europeans (see below) more helpful than the works of Townsend for understanding the major groupings of Tachinidae. Although this led Thompson to classify the Tachinidae of Trinidad in a more natural way, he had a proclivity for describing unnecessary new genera.

-

The tribes, genera and species created by Townsend were described predominantly for New World taxa and by their sheer number continue to pose serious challenges for taxonomists to this day. Sabrosky and Arnaud (1965), the first to catalogue the Tachinidae of America north of Mexico in the post-Townsend era, adopted a nearly modern concept of the family (differing only by the inclusion of Rhinophorinae) while otherwise retaining many of Townsend’s tribes:

-

“for present convenience, in the absence of any other published arrangement of the Nearctic genera, though with some combinations and generic transfers, notably where we agree with the recent work of Mesnil and coworkers in Europe. This is especially true in the Goniinae [= Exoristinae]” (p. 962).

-

The catalogue of the Tachinidae of America south of the United States by Guimarães (1971) followed shortly after Thompson’s Tachinids of Trinidad and Sabrosky and Arnaud’s catalogue. This author, faced with the huge number of tribes, genera and species described by Townsend and having to deal with other taxa inadequately described by earlier authors, could not revise the whole classification and mostly followed Townsend. This action, he admitted, resulted in a “catalogue arrangement [that] leaves much to be desired” (Guimarães 1971: 3). The rich fauna of the region was catalogued into 2864 species and (by Guimarães’ own admission) an over-split 944 genera.

-

There have been to date only two major attempts to correct the generic imbalance that has impeded study of New World Tachinidae, both by Donald Montgomery Wood. The first was a conspectus of the Blondeliini of North and Central America and the West Indies (Wood 1985). Although this study excluded South American Blondeliini, it nevertheless reduced the number of valid genera from about 230 to 55. Many of the genera sunk into synonymy were Townsend’s but there were also many described by Reinhard, Thompson, Curran and others. The second work to reduce the number of New World genera was Wood’s (1987) Tachinidae chapter in Manual of Nearctic Diptera. The nomenclatural changes in this work, including almost 200 new generic synonyms, were later enumerated by O’Hara and Wood (1998).

-

Wood (1987) also successfully bridged the gap between the generic classifications of the Nearctic and Palaearctic regions created by Townsend some decades earlier. This was accomplished partly by reducing the number of genera but also by assessing genera from a Holarctic perspective. The catalogue by O’Hara and Wood (2004) further united the classifications of Nearctic and Palaearctic Tachinidae. The catalogue by Guimarães (1971) has not been updated and the 800+ genera currently recognized in America south of the United States will not be easily converted into a modern classification. A careful study of the name-bearing types of the type species of many of these genera will be necessary before a better classification can be constructed for Neotropical Tachinidae.

+

The tribes, genera and species created by Townsend were described predominantly for New World taxa and by their sheer number continue to pose serious challenges for taxonomists to this day. Sabrosky and Arnaud (1965), the first to catalogue the Tachinidae of America north of Mexico in the post-Townsend era, adopted a nearly modern concept of the family (differing only by the inclusion of Rhinophorinae) while otherwise retaining many of Townsend’s tribes:

+

“for present convenience, in the absence of any other published arrangement of the Nearctic genera, though with some combinations and generic transfers, notably where we agree with the recent work of Mesnil and coworkers in Europe. This is especially true in the Goniinae [= Exoristinae]” (p. 962).

+

The catalogue of the Tachinidae of America south of the United States by Guimarães (1971) followed shortly after Thompson’s Tachinids of Trinidad and Sabrosky and Arnaud’s catalogue. This author, faced with the huge number of tribes, genera and species described by Townsend and having to deal with other taxa inadequately described by earlier authors, could not revise the whole classification and mostly followed Townsend. This action, he admitted, resulted in a “catalogue arrangement [that] leaves much to be desired” (Guimarães 1971: 3). The rich fauna of the region was catalogued into 2864 species and (by Guimarães’ own admission) an over-split 944 genera.

+

There have been to date only two major attempts to correct the generic imbalance that has impeded study of New World Tachinidae, both by Donald Montgomery Wood. The first was a conspectus of the Blondeliini of North and Central America and the West Indies (Wood 1985). Although this study excluded South American Blondeliini, it nevertheless reduced the number of valid genera from about 230 to 55. Many of the genera sunk into synonymy were Townsend’s but there were also many described by Reinhard, Thompson, Curran and others. The second work to reduce the number of New World genera was Wood’s (1987) Tachinidae chapter in Manual of Nearctic Diptera. The nomenclatural changes in this work, including almost 200 new generic synonyms, were later enumerated by O’Hara and Wood (1998).

+

Wood (1987) also successfully bridged the gap between the generic classifications of the Nearctic and Palaearctic regions created by Townsend some decades earlier. This was accomplished partly by reducing the number of genera but also by assessing genera from a Holarctic perspective. The catalogue by O’Hara and Wood (2004) further united the classifications of Nearctic and Palaearctic Tachinidae. The catalogue by Guimarães (1971) has not been updated and the 800+ genera currently recognized in America south of the United States will not be easily converted into a modern classification. A careful study of the name-bearing types of the type species of many of these genera will be necessary before a better classification can be constructed for Neotropical Tachinidae.

 The European influence -

The Europeans of the early 1900s continued to build on the discoveries of Girschner and others at the same time that Townsend in the New World was pursuing his own course of investigations that would culminate in his Manual of Myiology. Joseph Villeneuve de Janti, a medical doctor by profession (like Robineau-Desvoidy), emerged as an early specialist on the Tachinidae and published actively on the family from 1900 until his death in 1944. He wrote an influential paper in 1924 reviewing earlier works on chaetotaxy and detailing his own views on characters useful for understanding the evolution of the “Myodaires supérieurs”. This group comprised the “Tachinaires” (Tachinidae sensu present-day Oestroidea) and “Anthomyaires” (Anthomyiidae sensu present-day Muscoidea). Within Villeneuve’s Tachinidae were Calliphorinae, Sarcophaginae, Dexiinae, Rhinophorinae, Phasiinae, and Tachininae. Particularly noteworthy and progressive was the division of the Tachininae into two groups, Eutachininae and Protachininae, of which the former was considered more evolved than the latter. As a rough approximation, the two correspond to present-day Exoristinae and Tachininae, respectively.

+

The Europeans of the early 1900s continued to build on the discoveries of Girschner and others at the same time that Townsend in the New World was pursuing his own course of investigations that would culminate in his Manual of Myiology. Joseph Villeneuve de Janti, a medical doctor by profession (like Robineau-Desvoidy), emerged as an early specialist on the Tachinidae and published actively on the family from 1900 until his death in 1944. He wrote an influential paper in 1924 reviewing earlier works on chaetotaxy and detailing his own views on characters useful for understanding the evolution of the “Myodaires supérieurs”. This group comprised the “Tachinaires” (Tachinidae sensu present-day Oestroidea) and “Anthomyaires” (Anthomyiidae sensu present-day Muscoidea). Within Villeneuve’s Tachinidae were Calliphorinae, Sarcophaginae, Dexiinae, Rhinophorinae, Phasiinae, and Tachininae. Particularly noteworthy and progressive was the division of the Tachininae into two groups, Eutachininae and Protachininae, of which the former was considered more evolved than the latter. As a rough approximation, the two correspond to present-day Exoristinae and Tachininae, respectively.

Villeneuve was well respected by contemporaries for his expertise in Tachinidae and willingly shared his knowledge with others. As noted by Wainwright (1928: 141), Villeneuve:

-

“has contributed largely towards the reduction to something like order of our knowledge of these insects. Possibly the full value of his services to science may never be appreciated, because so many of the fruits of his labours have been given to the world by other workers, whom he has unselfishly and ungrudgingly assisted”.

-

The discovery by Malloch (1923) that the Tachinidae and Dexiidae can be distinguished from Sarcophagidae, Calliphoridae and Muscidae by an enlarged “metanotum” (subscutellum) was a highly significant development in the classification and differentiation of these flies. It was likely this discovery that led Villeneuve (1933) to revise his earlier classification and divide the “Tachinaires” into three groups:

+

“has contributed largely towards the reduction to something like order of our knowledge of these insects. Possibly the full value of his services to science may never be appreciated, because so many of the fruits of his labours have been given to the world by other workers, whom he has unselfishly and ungrudgingly assisted”.

+

The discovery by Malloch (1923) that the Tachinidae and Dexiidae can be distinguished from Sarcophagidae, Calliphoridae and Muscidae by an enlarged “metanotum” (subscutellum) was a highly significant development in the classification and differentiation of these flies. It was likely this discovery that led Villeneuve (1933) to revise his earlier classification and divide the “Tachinaires” into three groups:

1) Tachinidae with Phasiinae, Dexiinae and Tachininae,

2) Sarcophagidae with Miltogramminae, Sarcophaginae and Calliphorinae, and

3) Rhinophoridae, a small group of isopod parasitoids.

Villeneuve (1933) treated the Eutachininae and Protachininae of the Tachininae at length.

Villeneuve was a mentor and friend of Louis Paul Mesnil, who was 36 years his junior (Mesnil 1950). It was originally Villeneuve who was invited by Lindner to author the Tachinidae volumes of his ambitious Die Fliegen der palaearktischen Region (hereafter FPR). However, as the project drew closer Villeneuve realized that the talented and younger Mesnil was a better choice to take on this demanding and long-term task (Herting 1987).

-

Mesnil was an avid student of Tachinidae. He demonstrated his enthusiasm and insight early by publishing, as one of his first works on the group, a lengthy treatise entitled Essai sur les Tachinaires (Mesnil 1939). He began the Essai by reviewing and critiquing the classifications of his more illustrious predecessors: Robineau-Desvoidy, Macquart, Meigen, Rondani, Brauer and Bergenstamm, Pandellé, and Girschner. In proposing a new classification, Mesnil (1939) drew special inspiration from the works of Robineau-Desvoidy and Villeneuve, and like Brauer and Bergenstamm, started by grouping together related genera and building the classification “depuis la base vers le sommet” [“from the base to the summit”] (p. 20).

-

Mesnil (1939) restricted the term Tachinaires to the family Larvaevoridae3 (i.e., Tachinidae). The main diagnostic feature of the family was the well-developed “postscutellum” (subscutellum), as previously implied in Villeneuve’s (1933) classification and explicitly adopted by Curran (1934, as “metanotum”). Mesnil relegated the Rhinophorinae and Sarcophaginae to the Calliphoridae and subdivided the Larvaevoridae into six subfamilies: Salmaciinae4, Phorocerinae, Larvaevorinae, Ameniinae, Dexiinae and Phasiinae (including Oestrini). These were keyed and characterized and most were further subdivided.

-

The Phorocerinae of Mesnil (1939) consisted of tachinids possessing a haired prosternum and a small “prealar” (postsutural supraalar) seta. Included within the Phorocerinae were three tribes: Phorocerini, Blondeliini, and Crocutini5. The Phorocerini, with vein M (as “4e”) having an angular bend and a shadow fold, and the Blondeliini, with vein M having a rounded bend and no shadow fold, and both possessing divergent subapical scutellar setae (convergent in Crocutini), have continued to the present virtually unchanged in their characterization (Wood 1972, 1985). The Phorocerini have since become known as the Exoristini.

-

Mesnil began publishing FPR instalments a few years after his Essai. The goal was to treat all of the Palaearctic Tachinidae to species level but the task proved too great for him alone. After 35 years and some 1500 pages of text, the Larvaevorinae (present-day Exoristinae and Tachininae) were completed (Mesnil 1944–1975) along with one instalment on the Dexiinae (Mesnil 1980). Herting planned to publish on the remainder of the Dexiinae and all of the Phasiinae but only one instalment on the latter was published (Herting 1983).

-

Mesnil’s (1944) first instalment for FPR began, as did his Essai, with general remarks about previous workers and their classifications. Mesnil (1944: 2) made these observations about the generic concepts of other workers:

-

“Oft auch haben sie alte künstliche Gattungen aufrechterhalten, deren Umfang jedes Maß überschreitet und deren Heterogenität offenkundig ist; können sie doch sogar Arten verschiedener Tribus enthalten.

+

Mesnil was an avid student of Tachinidae. He demonstrated his enthusiasm and insight early by publishing, as one of his first works on the group, a lengthy treatise entitled Essai sur les Tachinaires (Mesnil 1939). He began the Essai by reviewing and critiquing the classifications of his more illustrious predecessors: Robineau-Desvoidy, Macquart, Meigen, Rondani, Brauer and Bergenstamm, Pandellé, and Girschner. In proposing a new classification, Mesnil (1939) drew special inspiration from the works of Robineau-Desvoidy and Villeneuve, and like Brauer and Bergenstamm, started by grouping together related genera and building the classification “depuis la base vers le sommet” [“from the base to the summit”] (p. 20).

+

Mesnil (1939) restricted the term Tachinaires to the family Larvaevoridae3 (i.e., Tachinidae). The main diagnostic feature of the family was the well-developed “postscutellum” (subscutellum), as previously implied in Villeneuve’s (1933) classification and explicitly adopted by Curran (1934, as “metanotum”). Mesnil relegated the Rhinophorinae and Sarcophaginae to the Calliphoridae and subdivided the Larvaevoridae into six subfamilies: Salmaciinae4, Phorocerinae, Larvaevorinae, Ameniinae, Dexiinae and Phasiinae (including Oestrini). These were keyed and characterized and most were further subdivided.

+

The Phorocerinae of Mesnil (1939) consisted of tachinids possessing a haired prosternum and a small “prealar” (postsutural supraalar) seta. Included within the Phorocerinae were three tribes: Phorocerini, Blondeliini, and Crocutini5. The Phorocerini, with vein M (as “4e”) having an angular bend and a shadow fold, and the Blondeliini, with vein M having a rounded bend and no shadow fold, and both possessing divergent subapical scutellar setae (convergent in Crocutini), have continued to the present virtually unchanged in their characterization (Wood 1972, 1985). The Phorocerini have since become known as the Exoristini.

+

Mesnil began publishing FPR instalments a few years after his Essai. The goal was to treat all of the Palaearctic Tachinidae to species level but the task proved too great for him alone. After 35 years and some 1500 pages of text, the Larvaevorinae (present-day Exoristinae and Tachininae) were completed (Mesnil 1944–1975) along with one instalment on the Dexiinae (Mesnil 1980). Herting planned to publish on the remainder of the Dexiinae and all of the Phasiinae but only one instalment on the latter was published (Herting 1983).

+

Mesnil’s (1944) first instalment for FPR began, as did his Essai, with general remarks about previous workers and their classifications. Mesnil (1944: 2) made these observations about the generic concepts of other workers:

+

“Oft auch haben sie alte künstliche Gattungen aufrechterhalten, deren Umfang jedes Maß überschreitet und deren Heterogenität offenkundig ist; können sie doch sogar Arten verschiedener Tribus enthalten.

-

So lassen sich die meisten neuzeitlichen Dipterologen, da sie die wahren Merkmale der Tachinen zu wenig berücksichtigt haben, nach zwei Richtungen gruppieren: die einen unterteilen die Gattungen bis ins Unendliche und machen so fast alle monospezifisch (T. Townsend), die andern vereinigen zahlreiche Gattungen zu einem Ganzen und gelangen so zu monströsen Zusammenfassungen (Curran).”

-

[“Often, they have maintained old artificial genera whose scope exceeds all bounds and whose heterogeneity is obvious; even though they may contain species of different tribes.

-

Since most modern dipterists have not taken the true characteristics of tachinids into account, they can be grouped in two directions: some subdivide the genera into infinity and thus make almost all of them monospecific (T. Townsend), others unite numerous genera into a whole and arrive at monstrous compilations (Curran)”.]

-

Lindner (1933) established the classification of the Diptera that would be followed in FPR six years before Mesnil’s (1939) Essai. This constrained Mesnil (1944) into keeping Larvaevoridae in the older and broader sense of present-day Oestroidea instead of in the restricted sense of present-day Tachinidae. Recognized within Larvaevoridae were subfamilies Larvaevorinae (with tribes Salmaciini, Phorocerini and Larvaevorini), Dexiinae and Phasiinae. Mesnil’s (1939) Oestrini (then placed in Phasiinae) became the “Gastrophilinae” (Gasterophilinae), Oestrinae and “Hypoderminae” (Hypodermatinae) of Lindner (1933). It is clear that this higher classification did not appeal to Mesnil. To him, the true definition of the Larvaevoridae was undeniable (“unbestreitbare”) and based on the enlarged subscutellum and parasitic habits of the family (Mesnil 1944). His only recourse was to chart the classification he would have followed had he been permitted to do so (numbers in parentheses refer to Lindner’s numbering system for families) (Mesnil 1944: 20):

+

So lassen sich die meisten neuzeitlichen Dipterologen, da sie die wahren Merkmale der Tachinen zu wenig berücksichtigt haben, nach zwei Richtungen gruppieren: die einen unterteilen die Gattungen bis ins Unendliche und machen so fast alle monospezifisch (T. Townsend), die andern vereinigen zahlreiche Gattungen zu einem Ganzen und gelangen so zu monströsen Zusammenfassungen (Curran).”

+

[“Often, they have maintained old artificial genera whose scope exceeds all bounds and whose heterogeneity is obvious; even though they may contain species of different tribes.

+

Since most modern dipterists have not taken the true characteristics of tachinids into account, they can be grouped in two directions: some subdivide the genera into infinity and thus make almost all of them monospecific (T. Townsend), others unite numerous genera into a whole and arrive at monstrous compilations (Curran)”.]

+

Lindner (1933) established the classification of the Diptera that would be followed in FPR six years before Mesnil’s (1939) Essai. This constrained Mesnil (1944) into keeping Larvaevoridae in the older and broader sense of present-day Oestroidea instead of in the restricted sense of present-day Tachinidae. Recognized within Larvaevoridae were subfamilies Larvaevorinae (with tribes Salmaciini, Phorocerini and Larvaevorini), Dexiinae and Phasiinae. Mesnil’s (1939) Oestrini (then placed in Phasiinae) became the “Gastrophilinae” (Gasterophilinae), Oestrinae and “Hypoderminae” (Hypodermatinae) of Lindner (1933). It is clear that this higher classification did not appeal to Mesnil. To him, the true definition of the Larvaevoridae was undeniable (“unbestreitbare”) and based on the enlarged subscutellum and parasitic habits of the family (Mesnil 1944). His only recourse was to chart the classification he would have followed had he been permitted to do so (numbers in parentheses refer to Lindner’s numbering system for families) (Mesnil 1944: 20):

I Haplostomata Frey

II Thecostomata Frey

A Muscidae (63)

@@ -154,63 +154,63 @@

a Phasiinae (incl. Oestrini) (64c)

b Dexiinae, Ameniinae (64f)

c Larvaevorinae (64g)

-

The Lindner series was published in small instalments (“Lieferungen”), the length of each being determined by the number of printed signatures used per instalment. Frequently an instalment would end in the middle of a description or in the middle of a key. This may have been cost-effective for the publisher but created havoc nomenclaturally. New generic names, for example, were often nomina nuda in one instalment and not made available until years later in another instalment. A great number of such nomenclatural issues as they pertain to the Tachinidae were dealt with by O’Hara (1996), Evenhuis and O’Hara (2008), and Evenhuis et al. (2008).

-

Mesnil’s FPR instalments by definition dealt primarily with the Palaearctic fauna but incorporated information on the taxa of other regions, except for the nearly impenetrable taxa of Neotropical Tachinidae. The result, in concert with a great many papers published by Mesnil outside FPR, was a classification for the bulk of the Tachinidae that could be hailed by contemporaries as a leap forward in the quest for a scheme reflecting the true relationships of the family. The suprageneric classification of Townsend (1934–1942) was largely ignored by Europeans who were making progress through their own investigations.

-

The first of Mesnil’s (1944) instalments in FPR gave only a glimpse of the classification that would follow. The Ameniinae were transferred to the Calliphoridae and kept as a subfamily, although the family itself is not currently considered monophyletic (e.g., Rognes 1997, Kutty et al. 2010). Mesnil’s (1944) three tribes of Larvaevorinae were split over the duration of FPR into a number of subtribes: nine in Salmaciini, six in Phorocerini and over 40 in Larvaevorini. The Larvaevorini were revisited by Mesnil (1966) and reclassified as Tachinini s. str. and Voriini. All of the subtribes of Mesnil (1944–1975) are now generally tribes and tribe Larvaevorini is present-day Tachininae. Many of the tribes continue to this day virtually unchanged whereas a few have undergone dramatic restructuring in the light of subsequent discoveries. The most significant changes resulted from research on the female postabdomen by Herting (1957) and male genitalia by Verbeke (1962a).

-

Benno Herting began his career on Tachinidae much the same way as did Mesnil (and even Robineau-Desvoidy) with an early publication based on original and extensive research (Herting 1957). It was a study of the female postabdomen and was based on the examination of about 500 species of calyptrate flies. Information about eggs and first instar larvae were taken into account but unlike Townsend’s studies the focus was more on the morphology of the terminal segments of the postabdomen than on the internal reproductive system. Herting (1957) used his findings to characterize the structural features of the female postabdomen throughout the families, subfamilies and lower groups of the Calyptratae. He tried to interpret these findings in a phylogenetic context and to adjust the classification accordingly.

-

Five subfamilies of the Tachinidae were recognized by Herting (1957): Echinomyiinae6, Dexiinae, Phasiinae, Ocypterinae7, and Eutachininae. At a gross level, Echinomyiinae corresponded to the Protachininae of Villeneuve (1924, 1933) and to the Larvaevorinae of Mesnil (1939; and later, Larvaevorini of Mesnil 1966–1975); Ocypterinae was formerly treated within Phasiinae by both Villeneuve (1924, 1933) and Mesnil (1939); and Eutachininae was proposed by Villeneuve (1924) and corresponded to the Salmaciinae (-ini) and Phorocerinae (-ini) of Mesnil (1939, 1944). Herting (1957) treated the Oestridae as a separate family.

-

Herting (1957) followed Villeneuve (1924, 1933) in using the subfamily name Eutachininae in his classification. He subdivided this subfamily into the Goniini and Eutachinini. He could not find reliable characters in the female postabdomen to separate these tribes and therefore chose to organize his discussion according to the reproductive habits of the species. Oviparous species were placed in the Eutachinini and distributed mostly between the Winthemia Robineau-Desvoidy group and Eutachina8 Brauer and Bergenstamm group. These were essentially the Winthemiina and Phorocerina that Mesnil (1944) had placed in tribes Salmaciini and Phorocerini, respectively. Ovolarviparous species grouped by Mesnil (1944) in the Blondeliina (tribe Phorocerini) were also assigned to the Eutachinini. The ovolarviparous Siphona Meigen group (Siphonina, tribe Phorocerini, of Mesnil 1944) was more clearly defined but its placement in Eutachinini or Goniini was not discussed. Similarly, the “Ethylla” (Ethilla) Robineau-Desvoidy group was included in Eutachininae but its further placement was not discussed. No members of the Acemyina (tribe Phorocerini) of Mesnil (1944) were studied by Herting (1957).

-

The composition of Herting’s (1957) Goniini consisted of species with two reproductive modes. One is quite specialized and involves the production of tiny (microtype) eggs that females oviposit on the food plants of hosts. These eggs hatch only after ingestion by a potential host. This sort of egg and the biology associated with it were already well known as a result of earlier studies (e.g., Sasaki 1887, Townsend 1908, 1911, Pantel 19109). The rest of Herting’s (1957) Goniini were mostly ovolarviparous species with a few oviparous species. This broad concept of the Goniini was essentially the Salmaciinae (-ini) of Mesnil (1939, 1944) without Ethyllina and Winthemiina.

+

The Lindner series was published in small instalments (“Lieferungen”), the length of each being determined by the number of printed signatures used per instalment. Frequently an instalment would end in the middle of a description or in the middle of a key. This may have been cost-effective for the publisher but created havoc nomenclaturally. New generic names, for example, were often nomina nuda in one instalment and not made available until years later in another instalment. A great number of such nomenclatural issues as they pertain to the Tachinidae were dealt with by O’Hara (1996), Evenhuis and O’Hara (2008), and Evenhuis et al. (2008).

+

Mesnil’s FPR instalments by definition dealt primarily with the Palaearctic fauna but incorporated information on the taxa of other regions, except for the nearly impenetrable taxa of Neotropical Tachinidae. The result, in concert with a great many papers published by Mesnil outside FPR, was a classification for the bulk of the Tachinidae that could be hailed by contemporaries as a leap forward in the quest for a scheme reflecting the true relationships of the family. The suprageneric classification of Townsend (1934–1942) was largely ignored by Europeans who were making progress through their own investigations.

+

The first of Mesnil’s (1944) instalments in FPR gave only a glimpse of the classification that would follow. The Ameniinae were transferred to the Calliphoridae and kept as a subfamily, although the family itself is not currently considered monophyletic (e.g., Rognes 1997, Kutty et al. 2010). Mesnil’s (1944) three tribes of Larvaevorinae were split over the duration of FPR into a number of subtribes: nine in Salmaciini, six in Phorocerini and over 40 in Larvaevorini. The Larvaevorini were revisited by Mesnil (1966) and reclassified as Tachinini s. str. and Voriini. All of the subtribes of Mesnil (1944–1975) are now generally tribes and tribe Larvaevorini is present-day Tachininae. Many of the tribes continue to this day virtually unchanged whereas a few have undergone dramatic restructuring in the light of subsequent discoveries. The most significant changes resulted from research on the female postabdomen by Herting (1957) and male genitalia by Verbeke (1962a).

+

Benno Herting began his career on Tachinidae much the same way as did Mesnil (and even Robineau-Desvoidy) with an early publication based on original and extensive research (Herting 1957). It was a study of the female postabdomen and was based on the examination of about 500 species of calyptrate flies. Information about eggs and first instar larvae were taken into account but unlike Townsend’s studies the focus was more on the morphology of the terminal segments of the postabdomen than on the internal reproductive system. Herting (1957) used his findings to characterize the structural features of the female postabdomen throughout the families, subfamilies and lower groups of the Calyptratae. He tried to interpret these findings in a phylogenetic context and to adjust the classification accordingly.

+

Five subfamilies of the Tachinidae were recognized by Herting (1957): Echinomyiinae6, Dexiinae, Phasiinae, Ocypterinae7, and Eutachininae. At a gross level, Echinomyiinae corresponded to the Protachininae of Villeneuve (1924, 1933) and to the Larvaevorinae of Mesnil (1939; and later, Larvaevorini of Mesnil 1966–1975); Ocypterinae was formerly treated within Phasiinae by both Villeneuve (1924, 1933) and Mesnil (1939); and Eutachininae was proposed by Villeneuve (1924) and corresponded to the Salmaciinae (-ini) and Phorocerinae (-ini) of Mesnil (1939, 1944). Herting (1957) treated the Oestridae as a separate family.

+

Herting (1957) followed Villeneuve (1924, 1933) in using the subfamily name Eutachininae in his classification. He subdivided this subfamily into the Goniini and Eutachinini. He could not find reliable characters in the female postabdomen to separate these tribes and therefore chose to organize his discussion according to the reproductive habits of the species. Oviparous species were placed in the Eutachinini and distributed mostly between the Winthemia Robineau-Desvoidy group and Eutachina8 Brauer and Bergenstamm group. These were essentially the Winthemiina and Phorocerina that Mesnil (1944) had placed in tribes Salmaciini and Phorocerini, respectively. Ovolarviparous species grouped by Mesnil (1944) in the Blondeliina (tribe Phorocerini) were also assigned to the Eutachinini. The ovolarviparous Siphona Meigen group (Siphonina, tribe Phorocerini, of Mesnil 1944) was more clearly defined but its placement in Eutachinini or Goniini was not discussed. Similarly, the “Ethylla” (Ethilla) Robineau-Desvoidy group was included in Eutachininae but its further placement was not discussed. No members of the Acemyina (tribe Phorocerini) of Mesnil (1944) were studied by Herting (1957).

+

The composition of Herting’s (1957) Goniini consisted of species with two reproductive modes. One is quite specialized and involves the production of tiny (microtype) eggs that females oviposit on the food plants of hosts. These eggs hatch only after ingestion by a potential host. This sort of egg and the biology associated with it were already well known as a result of earlier studies (e.g., Sasaki 1887, Townsend 1908, 1911, Pantel 19109). The rest of Herting’s (1957) Goniini were mostly ovolarviparous species with a few oviparous species. This broad concept of the Goniini was essentially the Salmaciinae (-ini) of Mesnil (1939, 1944) without Ethyllina and Winthemiina.

Herting (1957) introduced an important change to the placement of the Voriini. The members of this tribe had been included in the Protachininae of Villeneuve (1924, 1933) and the nearly equivalent Larvaevorinae of Mesnil (1939). Herting (1957) placed the tribe in the Dexiinae, bringing to three the number of Palaearctic tribes recognized in the subfamily: Dexiini, Voriini and Dufouriini. This move was supported by female postabdominal characters and by features of the male genitalia communicated to Herting by Verbeke (see below).

-

Mesnil (1956–1965) published on the Phorocerini in FPR over a ten-year period. He subdivided the tribe into subtribes Phorocerina, Blondeliina, Atylomyina, Neominthoina, Acemyina, and Siphonina, describing all the Palaearctic species and working in the same meticulous way that he had earlier for the Salmaciini (Mesnil 1944–1956). He had already revised the Old World Phorocerina (as Phorocerini) in a separate publication (Mesnil 1946) that he had probably begun before starting FPR. Mesnil (1956–1965) was halfway through the Phorocerini when Herting published his next great work on the Tachinidae, a monograph on the biology of the West Palaearctic species (Herting 1960). This work had a different focus from his earlier study but included a hierarchical arrangement of taxa that the former work had lacked. A clear classification was in evidence and although it was congruent in many respects with Mesnil’s it differed from it in some significant ways. Herting (1960) proposed a major restructuring of Mesnil’s Salmaciini (Mesnil 1944–1956) and Phorocerini (Mesnil 1956–1965). Both were united to form the Exoristinae10, consisting of a broadly defined Goniini (see above), Ethillini (Mesnil’s Ethyllina and Atylomyina), and the following tribes that corresponded to Mesnil’s remaining subtribes (except for the mixed and non-Palaearctic Neominthoina): Winthemiini, Exoristini (Mesnil’s Phorocerina), Blondeliini, Acemyiini, and Siphonini.

-

Herting’s (1960) Echinomyiinae included just three tribes: Echinomyiini, Leskiini and Microphthalmini. This work was published after Mesnil (1939) but before the FPR instalments on the same group (Mesnil 1966–1975, as “Larvaevorini oder Tachinini”). Mesnil (1939) had treated this group as the Larvaevorinae and noted that it was very close to Villeneuve’s (1933) Protachininae except for the exclusion of section Winthemia (placed by Mesnil in Salmaciinae [= Villeneuve’s Eutachininae], as Winthemiini). Mesnil’s (1939) Larvaevorinae had consisted of eight tribes11: Campylochaetini, Athryciini, Larvaevorini, Rhamphinini, Leskiini, Minthoini, Thelairini, and Macquartiini. This heterogeneous assemblage was considerably altered by Herting (1960): Larvaevorini and part of Macquartiini were placed in Echinomyiini; Campylochaetini, Athryciini, Thelairini and part of Macquartiini (i.e., the Phyllomyina) were moved to Voriini in the Dexiinae; Minthoini were included in Leskiini; and Rhamphinini were not treated but were later placed in Voriini by Herting (1984). The Microphthalmini of Herting (1960) were moved to the Tachininae from Mesnil’s (1939) section Dexiosomina (Dexiini, Dexiinae).

-

At the same time that Mesnil (1956–1965) was working through the Phorocerini using external characters and Herting (1957, 1960) was studying the female postabdomen, Jean Verbeke (1962a, 1962b12, 1963) was investigating tachinid male genitalia. Verbeke was communicating some of his findings to Herting before publishing them himself, thus contributing at least to Herting’s concept of the Dexiinae (see above). Verbeke (op. cit.) recognized within the complexity of the male genitalia a few general “types” associated with three structures. Firstly, the connection between the basiphallus and distiphallus is either “direct and non-mobile” (type I) or “indirect and mobile” (type II). Secondly, the distiphallus either lacks (POS [= Phasia, Ocyptera, Strongygaster] type) or possesses (DEG [= Dexia, Echinomyia, Gonia] type) longitudinal ventral microstructures. Thirdly, the “posterior paramere” (pregonite) has three types: type A, lobe-like and sensorial; type B, intermediate; and type C, strap-like and connective. These structural types do not form unique combinations and Verbeke (1963: 4) understood that “this repeated appearance of similar structures in different groups implicates a parallelism between the male genitalia of these groups”. Verbeke (1962a) concluded that the Tachinidae were best divided into six subfamilies: Phasiinae were characterized on the basis of a POS type distiphallus, whereas other Tachinidae have a DEG type distiphallus; Echinomyiinae (i.e., Tachininae) and Eutachininae (i.e., Exoristinae) have a type I connection between basiphallus and distiphallus; Dexiinae and Voriinae have a type II connection between basiphallus and distiphallus; and Dufouriinae with tribes Macquartiini and Dufourini, the former with a type I connection between the basiphallus and distiphallus and the latter with a type II connection but both tribes having a pregonite of type B. The subfamily Dufouriinae was clearly one of convenience and was not thought to be monophyletic. Verbeke (1963: 3) noted:

-

“Many other characters prove the intermediate situation of both tribes [intermediate between Dexiinae-Voriinae and Echinomyiinae-Eutachininae, see illustration in Verbeke (1962a: 147)] and for this reason we fused them into a new subfamily”.

-

Herting (1957, 1960) was aware of Verbeke’s studies on the male genitalia in advance of the publications on this subject (Verbeke 1962a, 1963) and was also familiar with the pioneering work on male genitalia by Rubtzov (1951). Herting (1957) discovered that features in the female postabdomen—and corroborated by evidence from the male postabdomen—supported a new concept of the Dexiinae. The Dexiini, Voriini and Dufouriini were brought together to form the Dexiinae. Although this classification differed from the one proposed later by Verbeke (1962a, 1963), it can be seen that Verbeke’s type II phallus and type C pregonite accurately defines Herting’s (1957, 1960) Dexiinae. This understanding of the subfamily continues to this day (e.g., Herting 1984, Tschorsnig 1985, Wood 1987, Tschorsnig and Richter 1998, O’Hara and Wood 2004, Cerretti 2010). Verbeke’s Macquartini, the other half of his Dufouriinae, was placed by Herting (op. cit.) in the Echinomyiinae but not retained as a tribe.

-

Mesnil (1966–1975) next published a series of instalments in FPR on the Larvaevorini, or Tachinini s. lat. In the first instalment, Mesnil (1966) introduced some changes to his earlier classification of the Larvaevorinae (i.e., Tachinidae). The classification proposed consisted of six tribes (equivalent to subfamilies of other authors): Phasiini, Exoristini, Goniini, Dexiini, Voriini, and Tachinini s. str. (see chart, Mesnil 1966: 882). The first three were characterized as producing planoconvex eggs and the last three as producing membranous eggs. Herting (1966) also noted this distinction in egg type between what he considered the two lineages of Tachinidae. Mesnil (1966) recognized the Phasiini as distinct based on the POS-type distiphallus of Verbeke (1962a) and the characteristic female postabdomen of Herting (1957). An unusual group that defies easy placement to this day, the Eutherina, were placed in the Voriinae by Verbeke (1962a) (based on male genitalia) and in the Phasiinae (-ini) by both Herting (1966) and Mesnil (1966) (based on egg type).

+

Mesnil (1956–1965) published on the Phorocerini in FPR over a ten-year period. He subdivided the tribe into subtribes Phorocerina, Blondeliina, Atylomyina, Neominthoina, Acemyina, and Siphonina, describing all the Palaearctic species and working in the same meticulous way that he had earlier for the Salmaciini (Mesnil 1944–1956). He had already revised the Old World Phorocerina (as Phorocerini) in a separate publication (Mesnil 1946) that he had probably begun before starting FPR. Mesnil (1956–1965) was halfway through the Phorocerini when Herting published his next great work on the Tachinidae, a monograph on the biology of the West Palaearctic species (Herting 1960). This work had a different focus from his earlier study but included a hierarchical arrangement of taxa that the former work had lacked. A clear classification was in evidence and although it was congruent in many respects with Mesnil’s it differed from it in some significant ways. Herting (1960) proposed a major restructuring of Mesnil’s Salmaciini (Mesnil 1944–1956) and Phorocerini (Mesnil 1956–1965). Both were united to form the Exoristinae10, consisting of a broadly defined Goniini (see above), Ethillini (Mesnil’s Ethyllina and Atylomyina), and the following tribes that corresponded to Mesnil’s remaining subtribes (except for the mixed and non-Palaearctic Neominthoina): Winthemiini, Exoristini (Mesnil’s Phorocerina), Blondeliini, Acemyiini, and Siphonini.

+

Herting’s (1960) Echinomyiinae included just three tribes: Echinomyiini, Leskiini and Microphthalmini. This work was published after Mesnil (1939) but before the FPR instalments on the same group (Mesnil 1966–1975, as “Larvaevorini oder Tachinini”). Mesnil (1939) had treated this group as the Larvaevorinae and noted that it was very close to Villeneuve’s (1933) Protachininae except for the exclusion of section Winthemia (placed by Mesnil in Salmaciinae [= Villeneuve’s Eutachininae], as Winthemiini). Mesnil’s (1939) Larvaevorinae had consisted of eight tribes11: Campylochaetini, Athryciini, Larvaevorini, Rhamphinini, Leskiini, Minthoini, Thelairini, and Macquartiini. This heterogeneous assemblage was considerably altered by Herting (1960): Larvaevorini and part of Macquartiini were placed in Echinomyiini; Campylochaetini, Athryciini, Thelairini and part of Macquartiini (i.e., the Phyllomyina) were moved to Voriini in the Dexiinae; Minthoini were included in Leskiini; and Rhamphinini were not treated but were later placed in Voriini by Herting (1984). The Microphthalmini of Herting (1960) were moved to the Tachininae from Mesnil’s (1939) section Dexiosomina (Dexiini, Dexiinae).

+

At the same time that Mesnil (1956–1965) was working through the Phorocerini using external characters and Herting (1957, 1960) was studying the female postabdomen, Jean Verbeke (1962a, 1962b12, 1963) was investigating tachinid male genitalia. Verbeke was communicating some of his findings to Herting before publishing them himself, thus contributing at least to Herting’s concept of the Dexiinae (see above). Verbeke (op. cit.) recognized within the complexity of the male genitalia a few general “types” associated with three structures. Firstly, the connection between the basiphallus and distiphallus is either “direct and non-mobile” (type I) or “indirect and mobile” (type II). Secondly, the distiphallus either lacks (POS [= Phasia, Ocyptera, Strongygaster] type) or possesses (DEG [= Dexia, Echinomyia, Gonia] type) longitudinal ventral microstructures. Thirdly, the “posterior paramere” (pregonite) has three types: type A, lobe-like and sensorial; type B, intermediate; and type C, strap-like and connective. These structural types do not form unique combinations and Verbeke (1963: 4) understood that “this repeated appearance of similar structures in different groups implicates a parallelism between the male genitalia of these groups”. Verbeke (1962a) concluded that the Tachinidae were best divided into six subfamilies: Phasiinae were characterized on the basis of a POS type distiphallus, whereas other Tachinidae have a DEG type distiphallus; Echinomyiinae (i.e., Tachininae) and Eutachininae (i.e., Exoristinae) have a type I connection between basiphallus and distiphallus; Dexiinae and Voriinae have a type II connection between basiphallus and distiphallus; and Dufouriinae with tribes Macquartiini and Dufourini, the former with a type I connection between the basiphallus and distiphallus and the latter with a type II connection but both tribes having a pregonite of type B. The subfamily Dufouriinae was clearly one of convenience and was not thought to be monophyletic. Verbeke (1963: 3) noted:

+

“Many other characters prove the intermediate situation of both tribes [intermediate between Dexiinae-Voriinae and Echinomyiinae-Eutachininae, see illustration in Verbeke (1962a: 147)] and for this reason we fused them into a new subfamily”.

+

Herting (1957, 1960) was aware of Verbeke’s studies on the male genitalia in advance of the publications on this subject (Verbeke 1962a, 1963) and was also familiar with the pioneering work on male genitalia by Rubtzov (1951). Herting (1957) discovered that features in the female postabdomen—and corroborated by evidence from the male postabdomen—supported a new concept of the Dexiinae. The Dexiini, Voriini and Dufouriini were brought together to form the Dexiinae. Although this classification differed from the one proposed later by Verbeke (1962a, 1963), it can be seen that Verbeke’s type II phallus and type C pregonite accurately defines Herting’s (1957, 1960) Dexiinae. This understanding of the subfamily continues to this day (e.g., Herting 1984, Tschorsnig 1985, Wood 1987, Tschorsnig and Richter 1998, O’Hara and Wood 2004, Cerretti 2010). Verbeke’s Macquartini, the other half of his Dufouriinae, was placed by Herting (op. cit.) in the Echinomyiinae but not retained as a tribe.

+

Mesnil (1966–1975) next published a series of instalments in FPR on the Larvaevorini, or Tachinini s. lat. In the first instalment, Mesnil (1966) introduced some changes to his earlier classification of the Larvaevorinae (i.e., Tachinidae). The classification proposed consisted of six tribes (equivalent to subfamilies of other authors): Phasiini, Exoristini, Goniini, Dexiini, Voriini, and Tachinini s. str. (see chart, Mesnil 1966: 882). The first three were characterized as producing planoconvex eggs and the last three as producing membranous eggs. Herting (1966) also noted this distinction in egg type between what he considered the two lineages of Tachinidae. Mesnil (1966) recognized the Phasiini as distinct based on the POS-type distiphallus of Verbeke (1962a) and the characteristic female postabdomen of Herting (1957). An unusual group that defies easy placement to this day, the Eutherina, were placed in the Voriinae by Verbeke (1962a) (based on male genitalia) and in the Phasiinae (-ini) by both Herting (1966) and Mesnil (1966) (based on egg type).

Mesnil (1966) was further influenced by Herting (1957) and Verbeke (1962a) to remove the voriines from the Larvaevorinae (-ini) of Mesnil (1939, 1944) and place them next to the dexiines. He kept the groups separate as Voriini and Dexiini rather than place them in the Dexiinae as did Herting (1957). The Dufouriinae of Verbeke (1962a) were split along similar lines to Herting (1957, 1960) with the Dufourini moved to Voriini as Dufouriina and Macquartiini kept in Tachinini s. str. (as Macquartiina) following Mesnil (1939). The original Dufouriina of Mesnil (1939) was a mixed group placed in Phasiini of Phasiinae and included such aberrant genera as Graphogaster Rondani and Rondaniooestrus Villeneuve in addition to Dufouria Robineau-Desvoidy and other typical dufouriines. Mesnil (1966) treated a more restricted Dufouriina in Voriini, placed Graphogaster in the small subtribe Graphogastrina in Tachinini s. str., and recognized Rondaniooestrus as sole member of Rondaniooestrina in Phasiini.

-

The Tachinini s. str. of Mesnil (1966) were split among about 30 subtribes. This tribe was equivalent to Mesnil’s (1939) Larvaevorinae and its eight tribes except for the removal of the voriines. In revising the earlier classification of Mesnil (1939) for FPR, Mesnil (1966) reduced his former tribes to subtribes and raised some former sections to tribes (especially among the Larvaevorini and Macquartiini of Mesnil 1939). This classification bears some resemblance to the groupings of Brauer and Bergenstamm (1889–1895) and Townsend (1934–1942) and reflected the uncertainty inherent in attempting to classify this heterogeneous and likely polyphyletic assemblage.

-

The Dexiosomina, treated in Dexiini of Dexiinae by Mesnil (1939), became part of Mesnil’s (1966–1975) Microphthalmina in Tachinini s. str.

-

Over 30 years elapsed between Mesnil’s (1944–1975) first and last FPR instalments on the Larvaevorinae. Mesnil (1975a, 1975b) included an Addenda and Corrigenda at the end of the Larvaevorini section in which he made corrections to earlier mistakes, added notes, and revised certain groups. His most significant change concerned the Goniinae (Salmaciini of Mesnil 1944–1956; i.e., present-day Exoristinae). This group had been based on external characters and needed revision to conform to the reproductive types discussed by Herting (1957, 1960). Mesnil (1975a: 1374) concluded:

-

“Nach Untersuchungen, die besonders durch B. Herting 1957 … über die Anatomie des Postabdomens der mikrooviparen Weibchen durchgeführt wurden, ist es möglich, die Gattungen der Goniinae in 2 Triben zu ordnen: die Goniini Rob.-Desv. (1830) mit mikrotypen Eiern und die Eryciini Rob.-Desv. (1830), die ovolarvipar oder ovipar sind.” [“According to studies that have been carried out especially by B. Herting 1957 … on the anatomy of the postabdomen of microoviparous females, it is possible to arrange the genera of Goniinae into two tribes: the Goniini Rob.-Desv. (1830) with microtype eggs and Eryciini Rob.-Desv. (1830), which are ovolarviparous or oviparous.”]

-

Goniini (s. str.) + Eryciini of Mesnil (1975a, 1975b) corresponded to Goniini (s. lat.) + Winthemiina + Ethillina of Herting (1960). Mesnil’s restriction of the Goniini to microovolarviparous tachinids was a key development in the classification of the Exoristinae. Herting (1984) would later remove the Winthemiina and Ethillina from Eryciini and treat them as tribes of Exoristinae, thereby creating a concept of Goniini s. str. + Eryciini equaling that of Herting’s (1960) Goniini.

-

The microovolarviparous tachinids had been recognized informally as a natural group within a broader Goniini since Herting’s (1957) study of the female postabdomen. A few years later Herting (1960) again grouped these tachinids as the “Mikroovipare Arten” within his broadly defined Goniini. Herting was known to be in favour of classifying the Goniini in a more restricted sense even before this was proposed by Mesnil (1975a). Very likely the idea was more his than Mesnil’s, although the two colleagues surely discussed the issue and may have influenced each other in how best to classify these tachinids. What is known is that Herting corresponded with others about his thoughts on this suprageneric complex prior to Mesnil (1975a) publishing on it. This is evident in Crosskey’s (1973b: 77) comments on the tribal classification he was adopting for Australian Goniinae (i.e., Exoristinae):

-

“Herting (personal communication) considers that the multifarious genera of the Goniini-Carceliini-Sturmiini-Eryciini complex should be aggregated into two tribes (for which the names Eryciini and Goniini would be nomenclaturally correct) according to whether they have an ovolarviparous or a microoviparous reproductive habit. Such a course has much to commend it insofar as it would probably reflect the real phylogeny more accurately than the present tribal system. But it is impossible to adopt such a system as yet for the Australian fauna, in which the reproductive habit of most of the genera remains unstudied.”

-

Thompson (1963), based on his own study of innumerable dissections, also recognized the microovolarviparous tachinids as a distinct group and devoted a separate part of Tachinids of Trinidad to the “goniines with microtype eggs”. Thompson (1963: 258) noted: “In the classification of Townsend, species producing microtype eggs are scattered through at least 14 tribes: Eriothrixini, Compsilurini, Phoroceratini, Phorinini, Actiini, Hyperecteinini, Frontinini, Goniini, Belvosiini, Harrisiini, Sturmiini, Lydellini, Phrynoini and Trypherini.”

+

The Tachinini s. str. of Mesnil (1966) were split among about 30 subtribes. This tribe was equivalent to Mesnil’s (1939) Larvaevorinae and its eight tribes except for the removal of the voriines. In revising the earlier classification of Mesnil (1939) for FPR, Mesnil (1966) reduced his former tribes to subtribes and raised some former sections to tribes (especially among the Larvaevorini and Macquartiini of Mesnil 1939). This classification bears some resemblance to the groupings of Brauer and Bergenstamm (1889–1895) and Townsend (1934–1942) and reflected the uncertainty inherent in attempting to classify this heterogeneous and likely polyphyletic assemblage.

+

The Dexiosomina, treated in Dexiini of Dexiinae by Mesnil (1939), became part of Mesnil’s (1966–1975) Microphthalmina in Tachinini s. str.

+

Over 30 years elapsed between Mesnil’s (1944–1975) first and last FPR instalments on the Larvaevorinae. Mesnil (1975a, 1975b) included an Addenda and Corrigenda at the end of the Larvaevorini section in which he made corrections to earlier mistakes, added notes, and revised certain groups. His most significant change concerned the Goniinae (Salmaciini of Mesnil 1944–1956; i.e., present-day Exoristinae). This group had been based on external characters and needed revision to conform to the reproductive types discussed by Herting (1957, 1960). Mesnil (1975a: 1374) concluded:

+

“Nach Untersuchungen, die besonders durch B. Herting 1957 … über die Anatomie des Postabdomens der mikrooviparen Weibchen durchgeführt wurden, ist es möglich, die Gattungen der Goniinae in 2 Triben zu ordnen: die Goniini Rob.-Desv. (1830) mit mikrotypen Eiern und die Eryciini Rob.-Desv. (1830), die ovolarvipar oder ovipar sind.” [“According to studies that have been carried out especially by B. Herting 1957 … on the anatomy of the postabdomen of microoviparous females, it is possible to arrange the genera of Goniinae into two tribes: the Goniini Rob.-Desv. (1830) with microtype eggs and Eryciini Rob.-Desv. (1830), which are ovolarviparous or oviparous.”]

+

Goniini (s. str.) + Eryciini of Mesnil (1975a, 1975b) corresponded to Goniini (s. lat.) + Winthemiina + Ethillina of Herting (1960). Mesnil’s restriction of the Goniini to microovolarviparous tachinids was a key development in the classification of the Exoristinae. Herting (1984) would later remove the Winthemiina and Ethillina from Eryciini and treat them as tribes of Exoristinae, thereby creating a concept of Goniini s. str. + Eryciini equaling that of Herting’s (1960) Goniini.

+

The microovolarviparous tachinids had been recognized informally as a natural group within a broader Goniini since Herting’s (1957) study of the female postabdomen. A few years later Herting (1960) again grouped these tachinids as the “Mikroovipare Arten” within his broadly defined Goniini. Herting was known to be in favour of classifying the Goniini in a more restricted sense even before this was proposed by Mesnil (1975a). Very likely the idea was more his than Mesnil’s, although the two colleagues surely discussed the issue and may have influenced each other in how best to classify these tachinids. What is known is that Herting corresponded with others about his thoughts on this suprageneric complex prior to Mesnil (1975a) publishing on it. This is evident in Crosskey’s (1973b: 77) comments on the tribal classification he was adopting for Australian Goniinae (i.e., Exoristinae):

+

“Herting (personal communication) considers that the multifarious genera of the Goniini-Carceliini-Sturmiini-Eryciini complex should be aggregated into two tribes (for which the names Eryciini and Goniini would be nomenclaturally correct) according to whether they have an ovolarviparous or a microoviparous reproductive habit. Such a course has much to commend it insofar as it would probably reflect the real phylogeny more accurately than the present tribal system. But it is impossible to adopt such a system as yet for the Australian fauna, in which the reproductive habit of most of the genera remains unstudied.”

+

Thompson (1963), based on his own study of innumerable dissections, also recognized the microovolarviparous tachinids as a distinct group and devoted a separate part of Tachinids of Trinidad to the “goniines with microtype eggs”. Thompson (1963: 258) noted: “In the classification of Townsend, species producing microtype eggs are scattered through at least 14 tribes: Eriothrixini, Compsilurini, Phoroceratini, Phorinini, Actiini, Hyperecteinini, Frontinini, Goniini, Belvosiini, Harrisiini, Sturmiini, Lydellini, Phrynoini and Trypherini.”

Sabrosky and Arnaud (1965) (see also above) were caught between the Townsend legacy of New World tachinid taxonomy and the rapidly evolving views on tachinid relationships and classification of the European specialists Mesnil, Herting and Verbeke. Sabrosky and Arnaud (1965) recognized both the Goniini and Eryciini but neither tribe corresponded very closely to the Goniini and Eryciini later defined by Mesnil (1975a, 1975b).

-

There was no Palaearctic catalogue of Tachinidae published between those of Bezzi and Stein (1907) and Herting (1984). Authors in the Old World wishing to treat regional faunas during this period were given overviews of emerging classifications first by Villeneuve (1924, 1933) and then by Mesnil (1939, 1944–1975), with contributions in particular from Herting (1957, 1960) and Verbeke (1962a). Villeneuve was acknowledged as a significant influence in the regional treatments of Stein (1924), Lundbeck (1927) and Wainwright (1928). As noted above in a quote from Wainwright (1928), Villeneuve’s personal assistance to contemporary dipterists was as valuable a contribution to science as were his publications.

-

Before the Second World War, tachinid specimens from Africa were routinely sent to the Imperial (later Commonwealth) Institute of Entomology in London for identification, but in practise they were identified by Villeneuve in France. This changed when the war severed relations with Villeneuve and the task of identifying Tachinidae fell to the recently hired dipterist, Fritz Isidor van Emden. Thus began van Emden’s foray into the Tachinidae that resulted in his valuable contributions on the faunas of the Afrotropical (as “Ethiopian”) Region (van Emden 1945, 1947, 1960 [the last posthumously]) and British Isles (van Emden 1954). In choosing a classification to follow, van Emden (1954: 7) noted:

-

“a sound classification has only recently been suggested by Villeneuve (1924, 1933) and worked out by Mesnil (1939, 1944). Being of such recent date, this ingenious classification has not so far been checked and applied to the whole of the family.”

-

Van Emden was slightly too early to take advantage of the progress to come during the 1960s through the efforts of Mesnil, Herting and Verbeke. Van Emden had planned to prepare keys to the whole of the Afrotropical Tachinidae but died before the third part was published (van Emden 1960) and before the last and largest part (on Exoristinae, as “Goniinae”) could be started.

-

Dugdale (1969) was more fortunate in being able to consider the works of Herting (1957, 1960), Verbeke (1962a), and Dupuis (1963) along with the recently revised classification of Mesnil (1966) in his treatment of New Zealand Tachinidae. Dupuis (1963) had concerned himself exclusively with the Phasiinae and his classification of the subfamily differed from that of Verbeke’s principally in the exclusion of the Strongygasterini and Rondaniooestrini. Despite Dugdale’s (1969) detailed review of recent advances, the New Zealand fauna is a small and isolated one and the affinities of some of its taxa were not resolved by Dugdale and remain uncertain to this day.

+

There was no Palaearctic catalogue of Tachinidae published between those of Bezzi and Stein (1907) and Herting (1984). Authors in the Old World wishing to treat regional faunas during this period were given overviews of emerging classifications first by Villeneuve (1924, 1933) and then by Mesnil (1939, 1944–1975), with contributions in particular from Herting (1957, 1960) and Verbeke (1962a). Villeneuve was acknowledged as a significant influence in the regional treatments of Stein (1924), Lundbeck (1927) and Wainwright (1928). As noted above in a quote from Wainwright (1928), Villeneuve’s personal assistance to contemporary dipterists was as valuable a contribution to science as were his publications.

+

Before the Second World War, tachinid specimens from Africa were routinely sent to the Imperial (later Commonwealth) Institute of Entomology in London for identification, but in practise they were identified by Villeneuve in France. This changed when the war severed relations with Villeneuve and the task of identifying Tachinidae fell to the recently hired dipterist, Fritz Isidor van Emden. Thus began van Emden’s foray into the Tachinidae that resulted in his valuable contributions on the faunas of the Afrotropical (as “Ethiopian”) Region (van Emden 1945, 1947, 1960 [the last posthumously]) and British Isles (van Emden 1954). In choosing a classification to follow, van Emden (1954: 7) noted:

+

“a sound classification has only recently been suggested by Villeneuve (1924, 1933) and worked out by Mesnil (1939, 1944). Being of such recent date, this ingenious classification has not so far been checked and applied to the whole of the family.”

+

Van Emden was slightly too early to take advantage of the progress to come during the 1960s through the efforts of Mesnil, Herting and Verbeke. Van Emden had planned to prepare keys to the whole of the Afrotropical Tachinidae but died before the third part was published (van Emden 1960) and before the last and largest part (on Exoristinae, as “Goniinae”) could be started.

+

Dugdale (1969) was more fortunate in being able to consider the works of Herting (1957, 1960), Verbeke (1962a), and Dupuis (1963) along with the recently revised classification of Mesnil (1966) in his treatment of New Zealand Tachinidae. Dupuis (1963) had concerned himself exclusively with the Phasiinae and his classification of the subfamily differed from that of Verbeke’s principally in the exclusion of the Strongygasterini and Rondaniooestrini. Despite Dugdale’s (1969) detailed review of recent advances, the New Zealand fauna is a small and isolated one and the affinities of some of its taxa were not resolved by Dugdale and remain uncertain to this day.

Roger Ward Crosskey became the next dipterist with the Commonwealth Institute of Entomology after the death of van Emden. His would be a remarkable tenure, single-handedly producing a revision of the Rutiliini (a tribe of Dexiinae confined to the Oriental and Australasian regions, Crosskey 1973a), conspecti on the Tachinidae of Australia (Crosskey 1973b) and the Oriental Region (Crosskey 1976), a catalogue of the Afrotropical13 Tachinidae (Crosskey 1980b), and keys to the tachinid genera of tropical and southern Africa (Crosskey 1984). Additionally, Crosskey later assisted with the preparation of a catalogue of the Tachinidae of the Australasian and Oceanian regions (Cantrell and Crosskey 1989). These resources offered a wealth of information on the names, classification, identification and hosts of Old World non-Palaearctic Tachinidae. The function of these works, however, was not to investigate and further illuminate the phylogenetic relationships of the Tachinidae. Perhaps for this reason and for the sake of consistency, the classificatory scheme adopted for the earliest conspectus was carried through with little change to the final catalogue, despite advances in tachinid systematics in the interim.

-

The classifications of Crosskey (1973b, 1976, 1980b) and Cantrell and Crosskey (1989) are very nearly the same and are best compared to the overview of tachinid classification given by Mesnil (1966) and, with respect to the Goniini–Eryciini, Mesnil (1975a). The classification in these works differed from that of Mesnil most significantly in the following respects14:

-

1) Tachininae included, in addition to the Tachininae sensu Herting (1984), most of Mesnil’s (1966) Voriini as tribes Campylochetini, Parerigonini, Phyllomyini, Thelairini, Voriini, and Wagneriini. Mesnil’s (1966) voriine subtribe Dexiomimopsina was included in Leskiini (later, Dexiomimops Townsend was treated in Voriini of Dexiinae by Herting 1984 and Shima 1987).

-

2) The “Goniini-Carceliini-Sturmiini-Eryciini complex” of Goniinae (i.e., Exoristinae) was not divided into Goniini and Eryciini according to egg type as advocated by Herting (see quote above from Crosskey 1973b) and Mesnil (1975a). Crosskey (1973b) gave two practical reasons for this: the reproductive habits of most of the genera involved were unknown and separating the redefined Goniini and Eryciini in a key on the basis of external morphology would not be possible even if egg type of each genus was known.

+

The classifications of Crosskey (1973b, 1976, 1980b) and Cantrell and Crosskey (1989) are very nearly the same and are best compared to the overview of tachinid classification given by Mesnil (1966) and, with respect to the Goniini–Eryciini, Mesnil (1975a). The classification in these works differed from that of Mesnil most significantly in the following respects14:

+

1) Tachininae included, in addition to the Tachininae sensu Herting (1984), most of Mesnil’s (1966) Voriini as tribes Campylochetini, Parerigonini, Phyllomyini, Thelairini, Voriini, and Wagneriini. Mesnil’s (1966) voriine subtribe Dexiomimopsina was included in Leskiini (later, Dexiomimops Townsend was treated in Voriini of Dexiinae by Herting 1984 and Shima 1987).

+

2) The “Goniini-Carceliini-Sturmiini-Eryciini complex” of Goniinae (i.e., Exoristinae) was not divided into Goniini and Eryciini according to egg type as advocated by Herting (see quote above from Crosskey 1973b) and Mesnil (1975a). Crosskey (1973b) gave two practical reasons for this: the reproductive habits of most of the genera involved were unknown and separating the redefined Goniini and Eryciini in a key on the basis of external morphology would not be possible even if egg type of each genus was known.

3) Dufouriinae were recognized as a subfamily with tribes Dufouriini and Imitomyiini; Mesnil (1966) had treated the former as a subtribe of Voriini and the latter as a subtribe of Phasiini.

4) Doleschallini were recognized as a tribe of Dexiinae; Mesnil (1966) had treated the single Oriental/Australasian genus Doleschalla Walker in the Doleschallina of Voriini15.

5) Oxyphyllomyiini were recognized as a tribe of Tachininae; Mesnil (1966) had treated the single Oriental genus Oxyphyllomyia Villeneuve in the Oxyphyllomyiina of Voriini. Later, Shima (1983) transferred Oxyphyllomyia to Leskiini.

-

6) Thelairini of Tachininae included Mesnil’s (1966) Zambesina of Exoristini (see discussion, Crosskey 1973b: 75).

+

6) Thelairini of Tachininae included Mesnil’s (1966) Zambesina of Exoristini (see discussion, Crosskey 1973b: 75).

7) Palpostomatini and Glaurocarini were recognized as tribes of Tachininae; Mesnil (1966) had treated both as subtribes of Exoristini.

8) Neaerini and Siphonini were recognized as tribes of Exoristinae; Mesnil (1966) had treated both as subtribes of Tachinini.

9) Rondaniooestrini were placed in Tachininae; Mesnil (1966) had treated the Rondaniooestrina as a subtribe of Phasiini.

 The Modern Era -

Nearly 25 years after writing about the biology of the West Palaearctic Tachinidae (Herting 1960) and over 75 years after the Palaearctic Tachinidae were last catalogued (Bezzi and Stein 1907), Herting (1984) published a long-awaited Catalogue of Palearctic Tachinidae. Much had changed since the former catalogue, both in terms of the suprageneric classification and number of genera and species. The tachinid fauna of the Palaearctic Region was the most intensively studied of all regional faunas and an up-to-date catalogue was an invaluable resource. Mesnil’s classification had evolved significantly over the years since publication of Essai sur les Tachinaires in 1939 but the changes had taken place in stages and must not have been easy for a non-specialist to follow. Herting had introduced changes too, some accepted by Mesnil and others not. Coincidently, Herting’s (1984) catalogue came out at the end of Mesnil’s long career and there have not been any sweeping changes to tachinid classification since. What has changed will be discussed further on. Herting (1984: 2) compared his classification to that of Herting (1960):

-

“The subdivision into four subfamilies is the same, only the name Echinomyiinae had to be changed into Tachininae. Some alterations have been made on the tribal level: The tribe Goniini is now restricted to the microoviparous forms, whereas the oviparous and ovolarviparous genera are assembled in a separate tribe, Eryciini. In the subfamily Tachininae, the number of tribes has been moderately increased, but not all the divisions made by Mesnil (1966b) in Lindner 64g: 885–896, have been accepted. The Siphonini are transferred from the Exoristinae to the Tachininae, where they are certainly better placed.”

+

Nearly 25 years after writing about the biology of the West Palaearctic Tachinidae (Herting 1960) and over 75 years after the Palaearctic Tachinidae were last catalogued (Bezzi and Stein 1907), Herting (1984) published a long-awaited Catalogue of Palearctic Tachinidae. Much had changed since the former catalogue, both in terms of the suprageneric classification and number of genera and species. The tachinid fauna of the Palaearctic Region was the most intensively studied of all regional faunas and an up-to-date catalogue was an invaluable resource. Mesnil’s classification had evolved significantly over the years since publication of Essai sur les Tachinaires in 1939 but the changes had taken place in stages and must not have been easy for a non-specialist to follow. Herting had introduced changes too, some accepted by Mesnil and others not. Coincidently, Herting’s (1984) catalogue came out at the end of Mesnil’s long career and there have not been any sweeping changes to tachinid classification since. What has changed will be discussed further on. Herting (1984: 2) compared his classification to that of Herting (1960):

+

“The subdivision into four subfamilies is the same, only the name Echinomyiinae had to be changed into Tachininae. Some alterations have been made on the tribal level: The tribe Goniini is now restricted to the microoviparous forms, whereas the oviparous and ovolarviparous genera are assembled in a separate tribe, Eryciini. In the subfamily Tachininae, the number of tribes has been moderately increased, but not all the divisions made by Mesnil (1966b) in Lindner 64g: 885–896, have been accepted. The Siphonini are transferred from the Exoristinae to the Tachininae, where they are certainly better placed.”

The classification of Herting (1984) differed from that of Mesnil (1966, 1975a) primarily in the following respects:

1) Winthemiini and Ethillini were recognized as tribes of Exoristinae; Mesnil (1975a) had included them in Eryciini, the former as Winthemiina and the latter as the three subtribes Ethillina, Phorocerosomina, and Atylomyina.

2) Dufouriini were recognized as a tribe of Dexiinae; Mesnil (1966) had treated Dufouriina as a subtribe of Voriini.

-

3) Voriini were recognized, without subtribes, alongside Dexiini and Dufouriini as one of three Palaearctic tribes in Dexiinae. Mesnil (1966) had treated his Voriini on the same level as the Voriinae of Verbeke (1962a) with 17 subtribes (see above for the treatment of Mesnil’s Voriini in Tachininae by Crosskey).

+

3) Voriini were recognized, without subtribes, alongside Dexiini and Dufouriini as one of three Palaearctic tribes in Dexiinae. Mesnil (1966) had treated his Voriini on the same level as the Voriinae of Verbeke (1962a) with 17 subtribes (see above for the treatment of Mesnil’s Voriini in Tachininae by Crosskey).

4) Tribes of Tachininae were significantly reduced from the subtribes of Tachinini of Mesnil (1966), although there was a sizable increase over the three tribes formerly recognized by Herting (1960). This increase over Herting (1960) was due primarily to a finer splitting of Echinomyiini and the separation of Minthoini from Leskiini.

-

Following closely after Herting’s (1984) catalogue was a comprehensive and insightful study of the male postabdomen by Tschorsnig (1985), Herting’s student and later his successor in Stuttgart. Tschorsnig took a comparative approach, describing the structures comprising the male postabdomen, detailing variation throughout the family, and discussing at the end of each taxonomic group the evidence regarding affinities. The work was less focused on the phallus and the pre- and postgonites than that of Verbeke (1962a) and arrived at some different conclusions. For example, the Phasiinae were considered monophyletic based on the structure of the hypandrium rather than on Verbeke’s POS type distiphallus, and the Dexiinae of Herting (1984) and not Verbeke (1962a) were considered monophyletic based on Verbeke’s type II phallus and type C pregonite. Although Tschorsnig’s study was phylogenetic in nature it did not include a cladogram of inferred relationships. The author may have considered the subject too complex and uncertain to condense into a single cladogram and may have preferred instead to present information about possible relationships in a narrative format.

+

Following closely after Herting’s (1984) catalogue was a comprehensive and insightful study of the male postabdomen by Tschorsnig (1985), Herting’s student and later his successor in Stuttgart. Tschorsnig took a comparative approach, describing the structures comprising the male postabdomen, detailing variation throughout the family, and discussing at the end of each taxonomic group the evidence regarding affinities. The work was less focused on the phallus and the pre- and postgonites than that of Verbeke (1962a) and arrived at some different conclusions. For example, the Phasiinae were considered monophyletic based on the structure of the hypandrium rather than on Verbeke’s POS type distiphallus, and the Dexiinae of Herting (1984) and not Verbeke (1962a) were considered monophyletic based on Verbeke’s type II phallus and type C pregonite. Although Tschorsnig’s study was phylogenetic in nature it did not include a cladogram of inferred relationships. The author may have considered the subject too complex and uncertain to condense into a single cladogram and may have preferred instead to present information about possible relationships in a narrative format.

Cantrell (1988) also conducted a comparative study, this one on the postabdomen of both sexes of Australian Tachinidae with descriptions of first instars and puparia. It was based on a thesis that was presumably completed prior to the publication of Tschorsnig (1985) because this work was not cited. The study provided a good overview as well as notes about each tribe of Australian Tachinidae.

-

Herting’s (1984) catalogue has been particularly influential to modern tachinidology because it summarized the current state of knowledge after a long period of change and has been followed subsequently by a period of relative stability. There have been highly significant works on Tachinidae published since 1984 but no revolutionary ideas have emerged about higher level relationships and classification. This is not to say that Herting’s classification is a true reflection of tachinid phylogeny, but rather it has changed little because the large groups that are least likely to be monophyletic (e.g., Eryciini, Tachininae, Voriini) have remained too little understood to permit their reclassification along phylogenetic lines.

-

Some major regional treatments and larger taxonomic works since Herting (1984) are reviewed below. There is still uncertainty about the proper placement of certain taxa among some of these works and in comparison with the major works during Mesnil’s era. These differences mostly concern smaller taxonomic units, often genera, and rather than discuss them below they are listed in Table 1.

+

Herting’s (1984) catalogue has been particularly influential to modern tachinidology because it summarized the current state of knowledge after a long period of change and has been followed subsequently by a period of relative stability. There have been highly significant works on Tachinidae published since 1984 but no revolutionary ideas have emerged about higher level relationships and classification. This is not to say that Herting’s classification is a true reflection of tachinid phylogeny, but rather it has changed little because the large groups that are least likely to be monophyletic (e.g., Eryciini, Tachininae, Voriini) have remained too little understood to permit their reclassification along phylogenetic lines.

+

Some major regional treatments and larger taxonomic works since Herting (1984) are reviewed below. There is still uncertainty about the proper placement of certain taxa among some of these works and in comparison with the major works during Mesnil’s era. These differences mostly concern smaller taxonomic units, often genera, and rather than discuss them below they are listed in Table 1.

The varied taxonomic placements of certain taxa of the Tachinidae by different authors are shown. Square brackets are used to indicate that a family-group name based on the taxon in question is given in the work but the taxon itself is not named in the work.<br/>

@@ -239,12 +239,12 @@ Exoristinae, Acemyiini Dexiinae, Voriini Dexiinae, Dufouriini - — - — + — + — Tachininae, Microphthalmini - — - — - — + — + — + — Phasiinae, Strongygastrini Dexiinae, Voriini Goniini s. lat.6 @@ -260,27 +260,27 @@ Voriinae, Euthera group Dufouriinae, Macquartiini2 Echinomyiinae1, Microphthalma group - — + — Dufouriinae, Macquartiini2 Phasiinae, Strongygastrini7 Phasiinae, Strongygastrini7 Dexiinae, Thelairini Goniini s. lat.6 Echinomyiinae1, Gymnocheta group - — + — Voriinae Sabrosky and Arnaud (1965) Goniinae3, Acemyini Tachininae, Campylochetini - — + — Phasiinae, Eutherini Phasiinae, Imitomyiini Proseninae, Dexillini - — + — [Phasiinae, Palpostomatini] - — + — Phasiinae, Strongygastrini Dexiinae, Thelairini Goniini (restricted)6 @@ -307,16 +307,16 @@ Voriini s. lat. - Guimarães (1971) + Guimarães (1971) [Goniinae, Acemyini] Tachininae, Campylochetini - — + — Phasiinae, Eutherini - — + — Proseninae, Dexillini - — - — - — + — + — + — Phasiinae, Strongygastrini Dexiinae, Thelairini Goniini (restricted)6 @@ -335,7 +335,7 @@ Tachininae, Oxyphyllomyiini Tachininae, Palpostomatini Tachininae, Rondaniooestrini - — + — Tachininae, Thelairini Goniini (restricted)6 Goniinae, Neaerini @@ -348,11 +348,11 @@ Dexiinae, Voriini Dexiinae, Dufouriini Phasiinae, Eutherini - — + — Tachininae, Microphthalmini - — - — - — + — + — + — Phasiinae, Strongygastrini Dexiinae, Voriini microtype Goniini @@ -364,13 +364,13 @@ Cantrell and Crosskey (1989) [Goniinae, Acemyini] Tachininae, Campylochetini - —5 + —5 Phasiinae, Eutherini - — + — Tachininae, Microphthalmini - — + — Tachininae, Palpostomatini - — + — Phasiinae, Strongygastrini Dexiinae, Voriini Goniini (restricted)6 @@ -380,20 +380,20 @@ Shima (1989) - — - — - — + — + — + — Dexiinae - — - — - — - — - — + — + — + — + — + — Phasiinae, Strongygastrini - — + — microtype Goniini - — - — + — + — Dexiinae, Voriini @@ -402,11 +402,11 @@ Dexiinae, Voriini Dexiinae, Voriini Phasiinae, Eutherini - — + — Tachininae, Microphthalmini - — + — Tachininae, Palpostomatini - — + — Phasiinae, Strongygastrini Dexiinae, Voriini microtype Goniini @@ -422,9 +422,9 @@ Phasiinae, Eutherini Phasiinae Imitomyiini Tachininae, Microphthalmini - — - — - + — + — + Phasiinae, Strongygastrini Dexiinae, Voriini microtype Goniini @@ -433,16 +433,16 @@ Dexiinae, Voriini - O’Hara and Wood (2004) + O’Hara and Wood (2004) Tachininae, Acemyini Dexiinae, Campylochetini Dexiinae, Dufouriini Dexiinae, Eutherini Dexiinae, Imitomyiini Tachininae, Megaprosopini - — + — Dexiinae, Palpostomatini - — + — Phasiinae, Strongygastrini Dexiinae, Voriini microtype Goniini @@ -451,7 +451,7 @@ Dexiinae, Voriini - O’Hara et al. (2009) + O’Hara et al. (2009) Tachininae, Acemyini Dexiinae, Campylochetini Dexiinae, Dufouriini @@ -460,7 +460,7 @@ [Tachininae, Megaprosopini] Tachininae, Leskiini Tachininae, Palpostomatini - — + — Tachininae, Strongygastrini Dexiinae, Thelairini microtype Goniini @@ -476,9 +476,9 @@ Dexiinae, Eutherini [Dexiinae, Imitomyiini] Tachininae, Megaprosopini - — - — - — + — + — + — Tachininae, Strongygastrini Dexiinae, Voriini microtype Goniini @@ -493,29 +493,29 @@

1 Eutachininae and Echinomyiinae were used in the sense of Exoristinae and Tachininae (or Larvaevorinae), respectively.

 -

2 Verbeke (1962a) was often unclear when applying his findings to a classification. With respect to Imitomyia and Palpostoma, these were part of Macquartiini (also as ‘Macquartiines’) in Part 1 but were treated as Imitomyiini and Palpostomatini in Part 2, although in the latter they were presumably still in a subordinate relationship with Macquartiini of Part 1.

+

2 Verbeke (1962a) was often unclear when applying his findings to a classification. With respect to Imitomyia and Palpostoma, these were part of Macquartiini (also as ‘Macquartiines’) in Part 1 but were treated as Imitomyiini and Palpostomatini in Part 2, although in the latter they were presumably still in a subordinate relationship with Macquartiini of Part 1.

3 The name Goniinae was changed to Exoristinae when the latter was determined to have priority.

 -

4 Mesnil (1966: 882) treated all Tachinidae as Tachininae and recognized six tribes, the equivalent of other author’s subfamilies.

+

4 Mesnil (1966: 882) treated all Tachinidae as Tachininae and recognized six tribes, the equivalent of other author’s subfamilies.

5 Cantrell and Burwell (2010) recognized the Dufouriini as a tribe of Dexiinae.

 -

6 “Goniini s. lat.” comprises both microtype and non-microtype taxa and “Goniini (restricted)” comprises only a portion of the microtype taxa.

+

6 “Goniini s. lat.” comprises both microtype and non-microtype taxa and “Goniini (restricted)” comprises only a portion of the microtype taxa.

 -

7 Verbeke (1963) was not certain about the phylogenetic position of Strongygaster and Rondaniooestrus and suggested they might be “intermediaries” between Phasiinae and Dufouriinae.

+

7 Verbeke (1963) was not certain about the phylogenetic position of Strongygaster and Rondaniooestrus and suggested they might be “intermediaries” between Phasiinae and Dufouriinae.

 -

Among the larger regional treatments of the 1980s were Cantrell’s (1984) study of Australian Phasiinae and Wood’s (1985) conspectus of the Blondeliini of North and Central America and the West Indies (the latter discussed above). The first modern key to the genera of Nearctic Tachinidae was published by Wood (1987) in Manual of Nearctic Diptera. The Siphonini of the world were revised at the generic level by O’Hara (1989). The Tachinidae of the Australasian and Oceanian regions were catalogued by Cantrell and Crosskey (1989), not only bringing Crosskey’s (1973b) conspectus of Australian Tachinidae up-to-date but cataloguing for the first time the non-Australian tachinids of the Australasian and Oceanian regions. Shima (1989) published a general paper on tachinids aimed at a Japanese audience; this work, unpretentious in nature, was remarkably detailed and presented the first cladogram of inferred relationships among the major (and controversial) tachinid lineages.

-

Other than the detailed study of the systematics of Australasian Dexiini by Barraclough (1992), the 1990s were dominated by European authors. Pape (1992) published on the phylogeny of the Tachinidae family-group, wherein the Tachinidae were inferred to form a monophyletic group (see also analysis by Pape and Arnaud 2001). Belshaw (1993) produced a handbook to the tachinids of the British Isles, replacing the earlier handbook by van Emden (1954). A new Palaearctic catalogue of the Tachinidae was published by Herting and Dely-Draskovits (1993) in the series Catalogue of Palaearctic Diptera, essentially reproducing the catalogue of Herting (1984) with corrected spellings to conform with nomenclatural rules and including long lists of nomina dubia not given in the earlier catalogue. Tschorsnig and Herting (1994) produced a valuable work on the identification, distribution and ecology of the tachinids of Central Europe. Mihályi (1986) published a comprehensive identification guide to tachinid genera and species of Hungary. The Siphonini of Europe were revised by Andersen (1996). The Tachinidae chapter of Manual of Palaearctic Diptera was authored by Tschorsnig and Richter (1998), the Palaearctic equivalent of Wood’s (1987) chapter in Manual of Nearctic Diptera. Chao et al. (1998) reviewed the Tachinidae of China in Flies of China, with keys to species and numerous illustrations of external features and male genitalia. The first-ever detailed study of the puparia and larval cephalopharyngeal skeletons of Tachinidae was published by Ziegler (1998). Ziegler, in his phylogenetic conclusions (pp. 192–194), proposed placing Glaurocarini within Ormiini s. lat. and placing Dufouria Robineau-Desvoidy (type genus of Dufouriini) and Rondania Robineau-Desvoidy within Voriini s. lat. The decade closed with Sabrosky’s (1999) posthumously published volume on family-group names in Diptera. This work was about 50 years in the making and will be an indispensable reference for decades to come. The Tachinidae with 429 entries dwarfs all other dipteran families.

-

Traditional taxonomic works of the 21st Century began with a revision of the Polideini of America north of Mexico by O’Hara (2002). There followed a large and well-illustrated work on the identification of Tachinidae of the Russian Far East by Richter (2004). That same year, O’Hara and Wood (2004) published a catalogue of the Tachinidae of America north of Mexico (discussed above). In this work the previous classification of Sabrosky and Arnaud (1965) was revised to conform more closely to the European model of Herting (1984). An interactive online resource to the Tachinidae of Europe was produced by Tschorsnig et al. (2004) as part of the Fauna Europaea project and continues to provide easy access to names and distributions. A catalogue of the Tachinidae of China by O’Hara et al. (2009) provided information on the names, types, distributions, and references of the approximately 1100 species known from this country. The Manual of Central American Diptera included a chapter on the Tachinidae by Wood and Zumbado (2010) in which 232 genera were reviewed, keyed, and illustrated (mostly with figures from Wood 1987), thereby forming a fine companion to Wood (1987). A Ph.D. thesis formed the nucleus of Cerretti’s (2010) two-volume work on the Tachinidae of Italy. This treatise provided a wealth of general information on tachinids in addition to generic descriptions and keys to species of Italian Tachinidae. Also included was an interactive key to the tachinid genera of the West Palaearctic Region using the program MOSCH, developed primarily by Cerretti. An online MOSCH key to the tachinid genera of the Palaearctic Region was made available recently by Cerretti et al. (2012a).

-

The first molecular studies devoted to the Tachinidae made their appearance early in the 21st Century. The Exoristinae were the subject of Stireman’s (2002) molecular study of genes 28S rRNA and EF-1α. The results were only partly congruent with evidence derived from morphology, most notably in not supporting the monophyly of the Goniini. A reappraisal of the same data using a Bayesian analysis (Stireman 2005) did not produce a convincing consensus tree, suggesting that the chosen genes may not be good for inferring tribal relationships within Tachinidae. In a more recent study of the Exoristinae by Tachi and Shima (2010), four genes (white, 18S, 28S and 16S rDNA) were studied. The results were similar in most respects to those of Stireman (2002, 2005), although monophyly of the Goniini was supported. Kutty et al. (2010) examined nine gene regions to infer relationships within the Calyptratae and especially the Oestroidea. In this study their Tachinidae were either monophyletic or not, depending upon the type of analysis performed. In general, these early molecular studies have shown promise and more sophisticated approaches in the future using combined morphological and molecular data sets are expected to yield more convincing results.

+

Among the larger regional treatments of the 1980s were Cantrell’s (1984) study of Australian Phasiinae and Wood’s (1985) conspectus of the Blondeliini of North and Central America and the West Indies (the latter discussed above). The first modern key to the genera of Nearctic Tachinidae was published by Wood (1987) in Manual of Nearctic Diptera. The Siphonini of the world were revised at the generic level by O’Hara (1989). The Tachinidae of the Australasian and Oceanian regions were catalogued by Cantrell and Crosskey (1989), not only bringing Crosskey’s (1973b) conspectus of Australian Tachinidae up-to-date but cataloguing for the first time the non-Australian tachinids of the Australasian and Oceanian regions. Shima (1989) published a general paper on tachinids aimed at a Japanese audience; this work, unpretentious in nature, was remarkably detailed and presented the first cladogram of inferred relationships among the major (and controversial) tachinid lineages.

+

Other than the detailed study of the systematics of Australasian Dexiini by Barraclough (1992), the 1990s were dominated by European authors. Pape (1992) published on the phylogeny of the Tachinidae family-group, wherein the Tachinidae were inferred to form a monophyletic group (see also analysis by Pape and Arnaud 2001). Belshaw (1993) produced a handbook to the tachinids of the British Isles, replacing the earlier handbook by van Emden (1954). A new Palaearctic catalogue of the Tachinidae was published by Herting and Dely-Draskovits (1993) in the series Catalogue of Palaearctic Diptera, essentially reproducing the catalogue of Herting (1984) with corrected spellings to conform with nomenclatural rules and including long lists of nomina dubia not given in the earlier catalogue. Tschorsnig and Herting (1994) produced a valuable work on the identification, distribution and ecology of the tachinids of Central Europe. Mihályi (1986) published a comprehensive identification guide to tachinid genera and species of Hungary. The Siphonini of Europe were revised by Andersen (1996). The Tachinidae chapter of Manual of Palaearctic Diptera was authored by Tschorsnig and Richter (1998), the Palaearctic equivalent of Wood’s (1987) chapter in Manual of Nearctic Diptera. Chao et al. (1998) reviewed the Tachinidae of China in Flies of China, with keys to species and numerous illustrations of external features and male genitalia. The first-ever detailed study of the puparia and larval cephalopharyngeal skeletons of Tachinidae was published by Ziegler (1998). Ziegler, in his phylogenetic conclusions (pp. 192–194), proposed placing Glaurocarini within Ormiini s. lat. and placing Dufouria Robineau-Desvoidy (type genus of Dufouriini) and Rondania Robineau-Desvoidy within Voriini s. lat. The decade closed with Sabrosky’s (1999) posthumously published volume on family-group names in Diptera. This work was about 50 years in the making and will be an indispensable reference for decades to come. The Tachinidae with 429 entries dwarfs all other dipteran families.

+

Traditional taxonomic works of the 21st Century began with a revision of the Polideini of America north of Mexico by O’Hara (2002). There followed a large and well-illustrated work on the identification of Tachinidae of the Russian Far East by Richter (2004). That same year, O’Hara and Wood (2004) published a catalogue of the Tachinidae of America north of Mexico (discussed above). In this work the previous classification of Sabrosky and Arnaud (1965) was revised to conform more closely to the European model of Herting (1984). An interactive online resource to the Tachinidae of Europe was produced by Tschorsnig et al. (2004) as part of the Fauna Europaea project and continues to provide easy access to names and distributions. A catalogue of the Tachinidae of China by O’Hara et al. (2009) provided information on the names, types, distributions, and references of the approximately 1100 species known from this country. The Manual of Central American Diptera included a chapter on the Tachinidae by Wood and Zumbado (2010) in which 232 genera were reviewed, keyed, and illustrated (mostly with figures from Wood 1987), thereby forming a fine companion to Wood (1987). A Ph.D. thesis formed the nucleus of Cerretti’s (2010) two-volume work on the Tachinidae of Italy. This treatise provided a wealth of general information on tachinids in addition to generic descriptions and keys to species of Italian Tachinidae. Also included was an interactive key to the tachinid genera of the West Palaearctic Region using the program MOSCH, developed primarily by Cerretti. An online MOSCH key to the tachinid genera of the Palaearctic Region was made available recently by Cerretti et al. (2012a).

+

The first molecular studies devoted to the Tachinidae made their appearance early in the 21st Century. The Exoristinae were the subject of Stireman’s (2002) molecular study of genes 28S rRNA and EF-1α. The results were only partly congruent with evidence derived from morphology, most notably in not supporting the monophyly of the Goniini. A reappraisal of the same data using a Bayesian analysis (Stireman 2005) did not produce a convincing consensus tree, suggesting that the chosen genes may not be good for inferring tribal relationships within Tachinidae. In a more recent study of the Exoristinae by Tachi and Shima (2010), four genes (white, 18S, 28S and 16S rDNA) were studied. The results were similar in most respects to those of Stireman (2002, 2005), although monophyly of the Goniini was supported. Kutty et al. (2010) examined nine gene regions to infer relationships within the Calyptratae and especially the Oestroidea. In this study their Tachinidae were either monophyletic or not, depending upon the type of analysis performed. In general, these early molecular studies have shown promise and more sophisticated approaches in the future using combined morphological and molecular data sets are expected to yield more convincing results.

@@ -526,7 +526,7 @@ Acknowledgements -

I would like to thank Monty and Grace Wood (Ottawa), Neal Evenhuis (Bernice Pauahi Bishop Museum, Honolulu, USA), and John Stireman (Wright State University, Dayton, USA) for reading the first draft of this paper and offering helpful suggestions for improvement. Monty’s personal recollections from 50 years of studying tachinids and interacting with fellow tachinidologists were especially insightful. I also wish to express my gratitude to reviewers Hans-Peter Tschorsnig (Staatliches Museum für Naturkunde, Stuttgart, Germany), Norman Woodley (Systematic Entomology Lab, USDA, Washington, USA) and Joachim Ziegler (Humboldt-Universitaet zu Berlin, Berlin, Germany), and ZooKeys editor Pierfilippo Cerretti (Centro Nazionale Biodiversità Forestale “Bosco Fontana”, Verona, Italy), for their thoughtful comments on the submitted manuscript.

+

I would like to thank Monty and Grace Wood (Ottawa), Neal Evenhuis (Bernice Pauahi Bishop Museum, Honolulu, USA), and John Stireman (Wright State University, Dayton, USA) for reading the first draft of this paper and offering helpful suggestions for improvement. Monty’s personal recollections from 50 years of studying tachinids and interacting with fellow tachinidologists were especially insightful. I also wish to express my gratitude to reviewers Hans-Peter Tschorsnig (Staatliches Museum für Naturkunde, Stuttgart, Germany), Norman Woodley (Systematic Entomology Lab, USDA, Washington, USA) and Joachim Ziegler (Humboldt-Universitaet zu Berlin, Berlin, Germany), and ZooKeys editor Pierfilippo Cerretti (Centro Nazionale Biodiversità Forestale “Bosco Fontana”, Verona, Italy), for their thoughtful comments on the submitted manuscript.

 @@ -554,10 +554,10 @@

8 Eutachina Brauer and Bergenstamm, 1889 is currently a junior synonym of Exorista Meigen, 1803.

 -

9 It was Pantel (1910) who coined the term “microtype” for these tiny ingestible eggs of goniines.

+

9 It was Pantel (1910) who coined the term “microtype” for these tiny ingestible eggs of goniines.

 -

10 Mesnil (1956–1965) had called this tribe “Phorocerini oder Exoristini”. Nomenclaturally, Herting’s (1960) use of the name Exoristinae was simply an elevation of Mesnil’s Phorocerini to a subfamily under an alternate name.

+

10 Mesnil (1956–1965) had called this tribe “Phorocerini oder Exoristini”. Nomenclaturally, Herting’s (1960) use of the name Exoristinae was simply an elevation of Mesnil’s Phorocerini to a subfamily under an alternate name.

11 Mesnil (1939) referred to names ending in -inae as tribes and names ending in -ini as subtribes. To avoid confusion within this paper such names are called subfamilies and tribes, respectively.

@@ -579,139 +579,139 @@ References Aldrich JM (1905) A catalogue of North American Diptera (or two-winged flies). Smithsonian Miscellaneous Collections 46 (2) [= No. 1444]: 680 pp. Andersen S (1996) The Siphonini (Diptera: Tachinidae) of Europe. Fauna Entomologica Scandinavica 33: 146 pp. - Arnaud PH Jr (1958) The entomological publications of Charles Henry Tyler Townsend [1863–1944]; with lists of his new generic and specific names. Microentomology 23: 1-63. - Barraclough DA (1992) The systematics of the Australasian Dexiini (Diptera: Tachinidae: Dexiinae) with revisions of endemic genera. Invertebrate Taxonomy 6: 1127-1371. + Arnaud PH Jr (1958) The entomological publications of Charles Henry Tyler Townsend [1863–1944]; with lists of his new generic and specific names. Microentomology 23: 1-63. + Barraclough DA (1992) The systematics of the Australasian Dexiini (Diptera: Tachinidae: Dexiinae) with revisions of endemic genera. Invertebrate Taxonomy 6: 1127-1371.10.1071/it9921127 Belshaw R (1993) Tachinid flies. Diptera: Tachinidae. Handbooks for the identification of British Insects, 10, Part 4a(i). Royal Entomological Society of London, London. 169 pp. - Bezzi MStein P (1907) Cyclorrapha Aschiza. Cyclorrapha Schizophora: Schizometopa. Pp. 1–747. In: Becker T, Bezzi M, Kertész K, Stein P (Eds) Katalog der paläarktischen Dipteren. Band III. Budapest. 828 pp. - Brauer FBergenstamm JE von (1889–1895) Die Zweiflügler des Kaiserlichen Museums zu Wien. IV–VII. Vorarbeiten zu einer Monographie der Muscaria Schizometopa (exclusive Anthomyidae). Partes I– IV. F Tempsky, Wien. + Bezzi MStein P (1907) Cyclorrapha Aschiza. Cyclorrapha Schizophora: Schizometopa. Pp. 1–747. In: Becker T, Bezzi M, Kertész K, Stein P (Eds) Katalog der paläarktischen Dipteren. Band III. Budapest. 828 pp. + Brauer FBergenstamm JE von (1889–1895) Die Zweiflügler des Kaiserlichen Museums zu Wien. IV–VII. Vorarbeiten zu einer Monographie der Muscaria Schizometopa (exclusive Anthomyidae). Partes I– IV. F Tempsky, Wien. Cantrell BK (1984) Synopsis of the Australian Phasiinae, including revisions of Gerocyptera Townsend and the Australian species of Cylindromyia Meigen (Diptera: Tachinidae). Australian Journal of Zoology. Supplementary Series 102: 1-60. doi: 10.1071/AJZS102 Cantrell BK (1988) The comparative morphology of the male and female postabdomen of the Australian Tachinidae (Diptera), with descriptions of some first-instar larvae and pupae. Invertebrate Taxonomy 2: 81-221. Cantrell BKBurwell CJ (2010) The tribe Dufouriini (Diptera: Tachinidae: Dexiinae) recorded from Australia with the description of two new species. Memoirs of the Queensland Museum 55: 119-133. Cantrell BKCrosskey RW (1989) Family Tachinidae. In: Evenhuis NL (Ed). Catalog of the Diptera of the Australasian and Oceanian regions. [Bishop Museum Special Publication 86.] Bishop Museum Press, EJ Brill: 733-784. 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Being a selected list of the books and prints of Diptera taxonomy from the beginning of Linnaean nomenclature to the end of the year 1930; containing information on the biographies, bibliographies, types, collections, and patronymic genera of the authors listed in this work; including detailed information on publication dates, original and subsequent editions, and other ancillary data concerning the publications listed herein. 2 volumes. Backhuys Publishers, Leiden. x + 871 pp. - Evenhuis NLO’Hara JE (2008) The status of Mesnil’s 1949 Die Fliegen genus-group names (Diptera: Tachinidae). Zootaxa 1827: 65-68. - Evenhuis NLO’Hara JEPape TPont AC (2010) Nomenclatural studies toward a world catalog of Diptera genus-group names. Part I. André-Jean-Baptiste Robineau-Desvoidy. Zootaxa 2373: 265 pp. + van Emden FI (1960) Keys to the Ethiopian Tachinidae—III. Macquartiinae. 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Zootaxa 2373: 265 pp. Evenhuis NLPape TPont AC (2008) The problems of subsequent typification in genus-group names and use of the Zoological Record: a study of selected post-1930 Diptera genus-group names without type species designations. Zootaxa 1912: 1-44. - Frey R (1921) Studien über den Bau des Mundes der niederen Diptera Schizophora nebst Bemerkungen über die Systematik dieser Dipterengruppe. Acta Societatis pro Fauna et Flora Fennica 48 (3): 245 + [2] pp. + 10 pls. + Frey R (1921) Studien über den Bau des Mundes der niederen Diptera Schizophora nebst Bemerkungen über die Systematik dieser Dipterengruppe. Acta Societatis pro Fauna et Flora Fennica 48 (3): 245 + [2] pp. + 10 pls. Girschner E (1893) Beitrag zur Systematik der Musciden. Berliner Entomologische Zeitschrift 38: 297-312. - Girschner E (1896) Ein neues Musciden-System auf Grund der Thoracalbeborstung und der Segmentierung des Hinterleibes. 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Townsend CHT (1911) Announcement of further results secured in the study of muscoid flies. Annals of the Entomological Society of America 4: 127-152. - Townsend CHT (1934) Manual of myiology in twelve parts. Part I. Development and structure. Privately published by Charles Townsend & Filhos, Itaquaquecetuba, São Paulo. 275 pp. - Townsend CHT (1934–1942) Manual of myiology in twelve parts. Privately published by Charles Townsend & Filhos, Itaquaquecetuba, São Paulo. [Each part individually paginated for a total of ca. 3760 pp.] + Townsend CHT (1934) Manual of myiology in twelve parts. Part I. Development and structure. Privately published by Charles Townsend & Filhos, Itaquaquecetuba, São Paulo. 275 pp. + Townsend CHT (1934–1942) Manual of myiology in twelve parts. Privately published by Charles Townsend & Filhos, Itaquaquecetuba, São Paulo. [Each part individually paginated for a total of ca. 3760 pp.] - Townsend CHT (1935) Manual of myiology in twelve parts. Part II. Muscoid classification and habits. Privately published by Charles Townsend & Filhos, Itaquaquecetuba, São Paulo. 289 pp. + 9 pls. + Townsend CHT (1935) Manual of myiology in twelve parts. Part II. Muscoid classification and habits. Privately published by Charles Townsend & Filhos, Itaquaquecetuba, São Paulo. 289 pp. + 9 pls. Townsend CHT (1943) Charles H.T. Townsend. Revista de Entomologia 14: 311-313. - Tschorsnig HP (1985) Taxonomie forstlich wichtiger Parasiten: Untersuchungen zur Struktur des männlichen Postabdomens der Raupenfliegen (Diptera, Tachinidae). Stuttgarter Beiträge zur Naturkunde. Serie A (Biologie) 383: 1-137. - Tschorsnig HPHerting B (1994) Die Raupenfliegen (Diptera: Tachinidae) Mitteleuropas: Bestimmungstabellen und Angaben zur Verbreitung und Ökologie der einzelnen Arten. Stuttgarter Beiträge zur Naturkunde. 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Wood DMZumbado MA (2010) Tachinidae (tachinid flies, parasitic flies). In: Brown BVBorkent ACumming JMWood DMWoodley NEZumbado MA (Eds). Manual of Central American Diptera. Volume 2. NRC Research Press, Ottawa: 1343-1417. - Wulp FM van der (1888–1903) Insecta. Diptera. In: Godman FD, Salvin O (Eds) Biologia Centrali-Americana, or, contributions to the knowledge of the fauna and flora of Mexico and Central America. Volume 2. Taylor & Francis, London. x + 489 pp. + 13 pls. + Wulp FM van der (1888–1903) Insecta. Diptera. In: Godman FD, Salvin O (Eds) Biologia Centrali-Americana, or, contributions to the knowledge of the fauna and flora of Mexico and Central America. Volume 2. Taylor & Francis, London. x + 489 pp. + 13 pls. Yeates DKWiegmann BMCourtney GWMeier RLambkin CPape T (2007) Phylogeny and systematics of Diptera: two decades of progress and prospects. Zootaxa 1668: 565-590. - Zetterstedt JW (1842–1860) Diptera Scandinaviae disposita et descripta. 14 volumes. 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Supplement 3: 244 pp. From 78059652f8f2e46c9a340a7ce87f31d2ae7b825e Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Wed, 17 Jul 2013 10:32:01 +0100 Subject: [PATCH 08/18] Result of running script to add DOIs to articles that lack them --- 10.3897_zookeys.316.5132.xml | 22 +++++++++++----------- 1 file changed, 11 insertions(+), 11 deletions(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index 51e4820..748a5fe 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -586,7 +586,7 @@ Brauer FBergenstamm JE von (1889–1895) Die Zweiflügler des Kaiserlichen Museums zu Wien. IV–VII. Vorarbeiten zu einer Monographie der Muscaria Schizometopa (exclusive Anthomyidae). Partes I– IV. F Tempsky, Wien. Cantrell BK (1984) Synopsis of the Australian Phasiinae, including revisions of Gerocyptera Townsend and the Australian species of Cylindromyia Meigen (Diptera: Tachinidae). Australian Journal of Zoology. Supplementary Series 102: 1-60. doi: 10.1071/AJZS102 - Cantrell BK (1988) The comparative morphology of the male and female postabdomen of the Australian Tachinidae (Diptera), with descriptions of some first-instar larvae and pupae. Invertebrate Taxonomy 2: 81-221. + Cantrell BK (1988) The comparative morphology of the male and female postabdomen of the Australian Tachinidae (Diptera), with descriptions of some first-instar larvae and pupae. Invertebrate Taxonomy 2: 81-221.10.1071/it9880081 Cantrell BKBurwell CJ (2010) The tribe Dufouriini (Diptera: Tachinidae: Dexiinae) recorded from Australia with the description of two new species. Memoirs of the Queensland Museum 55: 119-133. Cantrell BKCrosskey RW (1989) Family Tachinidae. In: Evenhuis NL (Ed). Catalog of the Diptera of the Australasian and Oceanian regions. [Bishop Museum Special Publication 86.] Bishop Museum Press, EJ Brill: 733-784. Cerretti P (2010) I tachinidi della fauna italiana (Diptera Tachinidae) con chiave interattiva dei generi ovest-paleartici. Cierre Edizioni, Verona. Volume I, 573 pp. Volume II, 339 pp. + CD-ROM. @@ -607,25 +607,25 @@ Emden FI van (1945) Keys to the Ethiopian Tachinidae.—I. Phasiinae. Proceedings of the Zoological Society of London 114 [1944]: 389–436 + 3 pls. Emden FI van (1947) Keys to the Ethiopian Tachinidae.—II. Dexiinae. Proceedings of the Zoological Society of London 116: 627–674 + 3 pls. Emden FI van (1954) Diptera Cyclorrhapha. Calyptrata (I). Section (a). Tachinidae and Calliphoridae. Handbooks for the identification of British insects, 10, Part 4(a). Royal Entomological Society of London, London. 133 pp. - van Emden FI (1960) Keys to the Ethiopian Tachinidae—III. Macquartiinae. Proceedings of the Zoological Society of London 134: 313-487. + van Emden FI (1960) Keys to the Ethiopian Tachinidae—III. Macquartiinae. Proceedings of the Zoological Society of London 134: 313-487.10.1111/j.1469-7998.1960.tb05596.x Evenhuis NL (1997) Litteratura taxonomica dipterorum (1758–1930). Being a selected list of the books and prints of Diptera taxonomy from the beginning of Linnaean nomenclature to the end of the year 1930; containing information on the biographies, bibliographies, types, collections, and patronymic genera of the authors listed in this work; including detailed information on publication dates, original and subsequent editions, and other ancillary data concerning the publications listed herein. 2 volumes. Backhuys Publishers, Leiden. x + 871 pp. Evenhuis NLO’Hara JE (2008) The status of Mesnil’s 1949 Die Fliegen genus-group names (Diptera: Tachinidae). Zootaxa 1827: 65-68. Evenhuis NLO’Hara JEPape TPont AC (2010) Nomenclatural studies toward a world catalog of Diptera genus-group names. Part I. André-Jean-Baptiste Robineau-Desvoidy. Zootaxa 2373: 265 pp. Evenhuis NLPape TPont AC (2008) The problems of subsequent typification in genus-group names and use of the Zoological Record: a study of selected post-1930 Diptera genus-group names without type species designations. Zootaxa 1912: 1-44. Frey R (1921) Studien über den Bau des Mundes der niederen Diptera Schizophora nebst Bemerkungen über die Systematik dieser Dipterengruppe. Acta Societatis pro Fauna et Flora Fennica 48 (3): 245 + [2] pp. + 10 pls. - Girschner E (1893) Beitrag zur Systematik der Musciden. Berliner Entomologische Zeitschrift 38: 297-312. + Girschner E (1893) Beitrag zur Systematik der Musciden. Berliner Entomologische Zeitschrift 38: 297-312.10.1002/mmnd.18940380305 Girschner E (1896) Ein neues Musciden-System auf Grund der Thoracalbeborstung und der Segmentierung des Hinterleibes. Jllustrierte Wochenschrift für Entomologie 1: 12–16, 30–32, 61–64, 105–112. Guimarães JH (1971) Family Tachinidae (Larvaevoridae). A Catalogue of the Diptera of the Americas South of the United States 104: 333 pp. - Herting B (1957) Das weibliche Postabdomen der calyptraten Fliegen (Diptera) und sein Merkmalswert für die Systematik der gruppe. Zeitschrift für Morphologie und Ökologie der Tiere 45: 429-461. + Herting B (1957) Das weibliche Postabdomen der calyptraten Fliegen (Diptera) und sein Merkmalswert für die Systematik der gruppe. Zeitschrift für Morphologie und Ökologie der Tiere 45: 429-461.10.1007/bf00389872 Herting B (1960) Biologie der westpaläarktischen Raupenfliegen (Dipt., Tachinidae). Monographien zur angewandten Entomologie 16: 188 pp. - Herting B (1966) Beiträge zur Kenntnis der europäischen Raupenfliegen (Dipt. Tachinidae). IX. Stuttgarter Beiträge zur Naturkunde 146: 1-12. + Herting B (1966) Beiträge zur Kenntnis der europäischen Raupenfliegen (Dipt. Tachinidae). IX. Stuttgarter Beiträge zur Naturkunde 146: 1-12.10.1007/bf02375740 Herting B (1983) 64c. Phasiinae. Die Fliegen der palaearktischen Region 9 (Lieferung 329): 1–88. Herting B (1984) Catalogue of Palearctic Tachinidae (Diptera). Stuttgarter Beiträge zur Naturkunde. Serie A (Biologie) 369: 228 pp. Herting B (1987) Louis Paul Mesnil. Entomologe. Jahreshefte der Gesellschaft für Naturkunde in Württemberg 142: 314-316. Herting BDely-Draskovits Á (1993) Family Tachinidae. In: Soós ÁPapp L (Eds). Catalogue of Palaearctic Diptera. Volume 13. Anthomyiidae—Tachinidae. Hungarian Natural History Museum, Budapest: 118-458. International Commission on Zoological Nomenclature (1963) Opinion 678. The suppression under the Plenary Powers of the pamphlet published by Meigen, 1800. Bulletin of Zoological Nomenclature 20: 339-342. - Kutty SNPape TWiegmann BMMeier R (2010) Molecular phylogeny of the Calyptratae (Diptera: Cyclorrhapha) with an emphasis on the superfamily Oestroidea and the position of Mystacinobiidae and McAlpine’s fly. Systematic Entomology 35: 614-635. + Kutty SNPape TWiegmann BMMeier R (2010) Molecular phylogeny of the Calyptratae (Diptera: Cyclorrhapha) with an emphasis on the superfamily Oestroidea and the position of Mystacinobiidae and McAlpine’s fly. Systematic Entomology 35: 614-635.10.1111/j.1365-3113.2010.00536.x Lindner E (1933) Handbuch. Die Fliegen der palaearktischen Region 1 (Lieferung 74): 161–208 + pl. XIII. [Dating and pagination based on Evenhuis (1997).] Lundbeck W (1927) Diptera Danica. Genera and species of flies hitherto found in Denmark. Part VII. Platypezidae, Tachinidae. GEC Gad, Copenhagen. 560 + [11 (Index)] pp. Macquart J (1838–1855) Diptères exotiques nouveaux ou peu connus. 10 parts (including supplements). NE Roret, Paris. [Additional to these 10 parts was another part that preceded them under the title Insectes diptères nouveaux ou peu connus, published by NE Roret in 1838.] @@ -648,12 +648,12 @@ Mesnil LP (1980) 64f. Dexiinae. Die Fliegen der palaearktischen Region 9 (Lieferung 323): 1–52. Mihályi F (1986) Diptera II. Tachinidae—Rhinophoridae. Fauna Hungariae 161: 425 pp. [In Hungarian] O’Hara JE (1989) Systematics of the genus group taxa of the Siphonini (Diptera: Tachinidae). Quaestiones Entomologicae 25: 1-229. - O’Hara JE (1996) The tachinid taxa of Louis P. Mesnil, with notes on nomenclature (Insecta: Diptera). Canadian Entomologist 128: 115-165. + O’Hara JE (1996) The tachinid taxa of Louis P. Mesnil, with notes on nomenclature (Insecta: Diptera). Canadian Entomologist 128: 115-165.10.4039/ent128115-1 O’Hara JE (2002) Revision of the Polideini (Tachinidae) of America north of Mexico. Studia Dipterologica. Supplement 10: 170 pp. O’Hara JE (2012) World genera of the Tachinidae (Diptera) and their regional occurrence. Version 7.0. PDF document, 75 pp. http://www.nadsdiptera.org/Tach/Genera/Gentach_ver7.pdf [accessed 3 February 2013] O’Hara JECerretti PPape TEvenhuis NL (2011) Nomenclatural studies toward a world list of Diptera genus-group names. Part II: Camillo Rondani. Zootaxa 3141: 1-268. O’Hara JEShima HZhang CT (2009) Annotated catalogue of the Tachinidae (Insecta: Diptera) of China. Zootaxa 2190: 1-236. - O’Hara JEWood DM (1998) Tachinidae (Diptera): nomenclatural review and changes, primarily for America north of Mexico. Canadian Entomologist 130: 751-774. + O’Hara JEWood DM (1998) Tachinidae (Diptera): nomenclatural review and changes, primarily for America north of Mexico. Canadian Entomologist 130: 751-774.10.4039/ent130751-6 O’Hara JEWood DM (2004) Catalogue of the Tachinidae (Diptera) of America north of Mexico. Memoirs on Entomology, International 18: iv + 410 pp. Osten-Sacken CR (1858) Catalogue of the described Diptera of North America. Smithsonian Miscellaneous Collections 3 (1) [= No. 102]: xx + 92 pp. Osten-Sacken CR (1878) Catalogue of the described Diptera of North America. Second edition. Smithsonian Miscellaneous Collections 26 (2) [= No. 270]: xlvii + [2] + 276 pp. @@ -667,7 +667,7 @@ Richter VA (2004) Fam. Tachinidae—tachinids. In: Sidorenko VS (Ed). Key to the insects of Russian Far East. Volume VI. Diptera and Siphonaptera. Part 3. Dal’nauka, Vladivostok: 148-398. [In Russian.] Robineau-Desvoidy JB (1830) Essai sur les myodaires. Mémoires présentés par divers Savans a l’Académie Royale des Sciences de l’Institut de France. Sciences Mathématiques et Physiques, Sér. 2, 2: 813 pp. Robineau-Desvoidy JB (1863) Histoire naturelle des diptères des environs de Paris. Oeuvre posthume du Dr Robineau-Desvoidy publiée par les soins de sa famille, sous la direction de M. H. Monceaux. V Masson et fils, Paris; F Wagner, Leipzig; and Williams & Norgate, London. Tome premier, xvi + 1143 pp. Tome second, 920 pp. - Rognes K (1997) The Calliphoridae (blowflies) (Diptera: Oestroidea) are not a monophyletic group. Cladistics 13: 27-66. + Rognes K (1997) The Calliphoridae (blowflies) (Diptera: Oestroidea) are not a monophyletic group. Cladistics 13: 27-66.10.1006/clad.1997.0031 Rondani C (1856–1880) Dipterologiae italicae prodromus. 7 volumes (comprising separate works and instalments in journals). [See Sabrosky (1961), Evenhuis (1997) and O’Hara et al. (2011) for the complicated publication details of this series.] Rubtzov IA (1951) [Contribution to the morphology and evolution of the abdomen and genitalia of phasiine flies (Diptera, Phasiidae s. l.). ] Trudy Vsesoyuznogo Entomologicheskogo Obshchestva 43: 171-249. [In Russian] Sabrosky CW (1961) Rondani’s “Dipterologiae italicae prodromus”. Annals of the Entomological Society of America 54: 827-831. @@ -680,10 +680,10 @@ Shima H (1987) A revision of the genus Dexiomimops Townsend (Diptera, Tachinidae). Sieboldia, Supplement 1987: 83-96. Shima H (1989) Parasitic way of life in tachinid flies. Insectarium 26: 4–9, 46–51, 88–94, 120–126. [In Japanese; cladogram on p. 125 reproduced with English caption in The Tachinid Times 3: 9 (1990).] Stein P (1924) Die verbreitetsten Tachiniden Mitteleuropas nach ihren Gattungen und Arten. Archiv für Naturgeschichte. Abteilung A 90 (6): 1-271. - Stireman JO III (2002) Phylogenetic relationships of tachinid flies in subfamily Exoristinae (Tachinidae: Diptera) based on 28S rDNA and elongation factor-1α. Systematic Entomology 27: 409-435. + Stireman JO III (2002) Phylogenetic relationships of tachinid flies in subfamily Exoristinae (Tachinidae: Diptera) based on 28S rDNA and elongation factor-1α. Systematic Entomology 27: 409-435.10.1046/j.1365-3113.2002.00187.x Stireman JO III (2005) Phylogenetic reconstruction of Exoristinae using molecular data: a Bayesian re-analysis. The Tachinid Times 18: 4-6. Stireman JO IIIO’Hara JEMoulton JKCerretti PWinkler IS (2013) Progress towards a phylogeny of world Tachinidae. Year 1. The Tachinid Times 26: 4-9. - Tachi TShima H (2010) Molecular phylogeny of the subfamily Exoristinae (Diptera, Tachinidae), with discussions on the evolutionary history of female oviposition strategy. Systematic Entomology 35: 148-163. + Tachi TShima H (2010) Molecular phylogeny of the subfamily Exoristinae (Diptera, Tachinidae), with discussions on the evolutionary history of female oviposition strategy. Systematic Entomology 35: 148-163.10.1111/j.1365-3113.2009.00497.x Thompson WR (1961) The tachinids (Diptera) of Trinidad. I. The voriines. Transactions of the American Entomological Society 87: 21–44 + 5 pls. Thompson WR (1961–1968) The tachinids of Trinidad. Parts I–VIII. [Parts published in five different journals.] Thompson WR (1963) The tachinids of Trinidad. III. The goniines with microtype eggs (Dipt. Tachinidae). Studia Entomologica 6: 257-404. From b234cd2c1827b96b578978e8ddcbabb971b2422e Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Wed, 17 Jul 2013 15:55:22 +0100 Subject: [PATCH 09/18] Replaced DOI 10.1006/clad.1997.0031 with DOI 10.1111/j.1096-0031.1997.tb00240.x See http://support.crossref.org/requests/17764 --- 10.3897_zookeys.316.5132.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.316.5132.xml b/10.3897_zookeys.316.5132.xml index 748a5fe..44bb589 100644 --- a/10.3897_zookeys.316.5132.xml +++ b/10.3897_zookeys.316.5132.xml @@ -667,7 +667,7 @@ Richter VA (2004) Fam. Tachinidae—tachinids. In: Sidorenko VS (Ed). Key to the insects of Russian Far East. Volume VI. Diptera and Siphonaptera. Part 3. Dal’nauka, Vladivostok: 148-398. [In Russian.] Robineau-Desvoidy JB (1830) Essai sur les myodaires. Mémoires présentés par divers Savans a l’Académie Royale des Sciences de l’Institut de France. Sciences Mathématiques et Physiques, Sér. 2, 2: 813 pp. Robineau-Desvoidy JB (1863) Histoire naturelle des diptères des environs de Paris. Oeuvre posthume du Dr Robineau-Desvoidy publiée par les soins de sa famille, sous la direction de M. H. Monceaux. V Masson et fils, Paris; F Wagner, Leipzig; and Williams & Norgate, London. Tome premier, xvi + 1143 pp. Tome second, 920 pp. - Rognes K (1997) The Calliphoridae (blowflies) (Diptera: Oestroidea) are not a monophyletic group. Cladistics 13: 27-66.10.1006/clad.1997.0031 + Rognes K (1997) The Calliphoridae (blowflies) (Diptera: Oestroidea) are not a monophyletic group. Cladistics 13: 27-66.10.1111/j.1096-0031.1997.tb00240.x Rondani C (1856–1880) Dipterologiae italicae prodromus. 7 volumes (comprising separate works and instalments in journals). [See Sabrosky (1961), Evenhuis (1997) and O’Hara et al. (2011) for the complicated publication details of this series.] Rubtzov IA (1951) [Contribution to the morphology and evolution of the abdomen and genitalia of phasiine flies (Diptera, Phasiidae s. l.). ] Trudy Vsesoyuznogo Entomologicheskogo Obshchestva 43: 171-249. [In Russian] Sabrosky CW (1961) Rondani’s “Dipterologiae italicae prodromus”. Annals of the Entomological Society of America 54: 827-831. From 723c29def341af1dea45a5a823bee37c013d3fcb Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Fri, 6 Sep 2013 10:19:02 +0100 Subject: [PATCH 10/18] B14 was not tagged correctly --- 10.3897_zookeys.183.3073.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.183.3073.xml b/10.3897_zookeys.183.3073.xml index 7b4c6b2..150862d 100644 --- a/10.3897_zookeys.183.3073.xml +++ b/10.3897_zookeys.183.3073.xml @@ -225,7 +225,7 @@ Chace FAJr (1972) The shrimps of the Smithsonian–Bredin Caribbean expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology 98: 1-179. Christoffersen ML (1979) Campagne de la Calypso au large des côtes Atlantiques de l’Amerique du Sud (1961–1962). I. Decapod Crustacea: Alpheoida. Annales de l’Institut Océanographique 55(Suppl.): 297–377. Christoffersen ML (1998) Malacostraca. Eucarida. 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Published by the author, Ithaca, NY. http://www.cladistics.com/about_winc.htm From 4a12fe23f987778bf3b5f976ba31d0fed72260b8 Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Wed, 16 Oct 2013 13:24:41 +0100 Subject: [PATCH 13/18] Edited B30 --- 10.3897_zookeys.310.4914.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.310.4914.xml b/10.3897_zookeys.310.4914.xml index 424d0f8..722f919 100644 --- a/10.3897_zookeys.310.4914.xml +++ b/10.3897_zookeys.310.4914.xml @@ -475,7 +475,7 @@ Papp L (1984) Family Thyreophoridae. Vol. 9. In: Soós Á, Papp L (Eds) Catalogue of Palearctic Diptera. Micropezidae-Agromyzidae. Akadémiai Kiadó, Budapest: 241−242. Rivosecchi L (2008) Aggiunte e correzioni alle checklist di alcune famiglie di Ditteri della fauna italiana (Diptera). Bollettino della Società entomologica italiana 140 (2): 95-103 - Robineau-Desvoidy JB (1830) Essai sur les myodaires. 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From 2a48fc651548684c68d02a7ff8a99f67a18d4416 Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 6 Jan 2014 16:56:45 +0000 Subject: [PATCH 14/18] Lots of corrections to references markup --- 10.3897_zookeys.211.3463.xml | 64 ++++++++++++++++++------------------ 1 file changed, 32 insertions(+), 32 deletions(-) diff --git a/10.3897_zookeys.211.3463.xml b/10.3897_zookeys.211.3463.xml index 17c6270..5a01fd6 100644 --- a/10.3897_zookeys.211.3463.xml +++ b/10.3897_zookeys.211.3463.xml @@ -1571,34 +1571,34 @@ Arnett RH (1947) Epiperipatus braziliensis (Bouvier) on Barro Colorado Island, Canal Zone. Entomological News 58: 59-60. Arnett RH (1961) The Onychophora of Jamaica. Entomological News 72: 213-220. Baehr M (1977) Über einige Onychophoren aus Australien und Tasmanien mit Beschreibung einer neuen Art und Anmerkung zur Stellung von Ooperipatus paradoxus Bouvier, 1915. 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Nature 46: 514. doi: 10.1038/046514a0 + Gravier CFage L (1925) Sur une nouvelle espéce de Péripate du Chili (Opisthopatus costesi). Annales des Sciences Naturelles, Zoologie et Biologie Animale [10e Série] 8: 185–200. Grimaldi DAEngel MSNascimbene PC (2002) Fossiliferous cretaceous amber from Myanmar (Burma): its rediscovery, biotic diversity, and paleontological significance. American Museum Novitates 3361: 3-72. doi: 10.1206/0003-0082(2002)361<0001:FCAFMB>2.0.CO;2 Grube E (1853) Über den Bau von Peripatus edwardsii. Müller’s Archives of Anatomy and Physiology [1853]: 322–360. Grube E (1866) Peripatus capensis. Jahresbericht der Schlesischen Gesellschaft für Vaterländische Cultur 43: 65-66. @@ -1717,8 +1717,8 @@ Ruhberg HMesibov RBriscoe DATait NN (1991) Tasmanipatus barretti gen. nov., sp. nov. and T. anophthalmus sp. nov.: two new and unusual onychophorans (Onychophora: Peripatopsidae) from northeastern Tasmania. Papers and Proceedings of the Royal Society of Tasmania 125: 7-10. Ruhberg HTait NNBriscoe DAStorch V (1988) Cephalofovea tomahmontis n. gen., n. sp., an Australian peripatopsid (Onychophora) with a unique cephalic pit. Zoologischer Anzeiger 221: 117-133. Sampaio-Costa CChagas-Junior ABaptista RLC (2009) Brazilian species of Onychophora with notes on their taxonomy and distribution. Zoologia 26: 553-561. - Sänger N (1871) Peripatus capensis Gr. and Peripatus Leuckartii n. sp.. Transactions of the Second Congress of Russian Naturalists in Moscow: 239–262 [in Russian] - Sänger H (1900) Peripatus capensis Sr. et Peripatus leuckartii, n. sp. Bulletin de la Société Philomathique de Paris [Serie 9] 3: 9–36. + Sänger N (1871) Peripatus capensis Gr. and Peripatus Leuckartii n. sp.. Transactions of the Second Congress of Russian Naturalists in Moscow: 239–262 [in Russian] + Sänger H (1900) Peripatus capensis Sr. et Peripatus leuckartii, n. sp. Bulletin de la Société Philomathique de Paris [Serie 9] 3: 9–36. Schmarda LK (1871) I. Ordnung. Malacopoda Blainville. Onychophora. Grube. In: Schmarda LK (Ed). Zoologie. Wilhelm Braumüller, Vienna, 1st ed., vol. 2: 74-76. Schmarda LK (1878) I. Ordnung. Malacopoda Blainville. Onychophora. Grube. In: Schmarda LK (Ed). Zoologie. Wilhelm Braumüller, Vienna, 2nd ed., vol. 2: 74-76. Sclater WL (1888) On the early stages of the development of South American species of Peripatus. Quarterly Journal of Microscopical Science 28: 343-363. @@ -1730,7 +1730,7 @@ Sherbon BJWalker MH (2004) A new species of Peripatopsis from South Africa, P. stelliporata, with observations on embryonic development and sperm degradation (Onychophora, Peripatopsidae). Journal of Zoology 264: 295-305. doi: 10.1017/S0952836904005783 Spencer B (1909) Description of a new species of Peripatoides from West Australia. Proceedings of the Royal Society of Victoria 21: 420-422. Stuhlmann F (1886) Die Reifung des Arthropodeneies nach Beobachtung an Insekten, Spinnen, Myriapoden und Peripatus. Berichte der Naturforschenden Gesellschaft zu Freiburg iB 1: 128. - Tait NNBriscoe DA (1995) Genetic differentiation within New Zealand Onychophora and their relationships to the Australian fauna. Zoological Journal of the Linnean Society 114: 103–113. doi: 10.1111/j.1096-3642.1995.tb00115.x + Tait NNBriscoe DA (1995) Genetic differentiation within New Zealand Onychophora and their relationships to the Australian fauna. Zoological Journal of the Linnean Society 114: 103–113. doi: 10.1111/j.1096-3642.1995.tb00115.x Tait NNNorman JM (2001) Novel mating behaviour in Florelliceps stutchburyae gen. nov., sp. nov. (Onychophora: Peripatopsidae) from Australia. Journal of Zoology 253: 301-308. Trewick SA (1998) Sympatric cryptic species in New Zealand Onychophora. Biological Journal of the Linnean Society 63: 307-329. doi: 10.1111/j.1095-8312.1998.tb01520.x Trindade G (1958) Peripatus e sua possível utilização em laboratório. Anais do III Congresso de Farmácia e Bioquímica Pan-americano e V Congresso Brasileiro de Farmácia: 519–520. @@ -1739,12 +1739,12 @@ Vasconcellos AAlmeida WOEloy ECC (2004) Onychophora de florestas úmidas do complexo da Mata Atlântica do nordeste brasileiro e sua importância para conservação e estudos sistemáticos. In: Pôrto KCCabral JJPTabarelli M (Eds). Brejos de Altitude: história natural, ecologia e conservação. Ministério do Meio Ambiente, Brasília: 139-144. von Kennel J (1883) Entwicklungsgeschichte von Peripatus. Zoologischer Anzeiger 6: 531-537. - Weber M (1898) Wetenschappelijke Vergadering. Tijdschrift der Nederlandsche Dierkundige Vereniging 5(4), VII-IX. + Weber M (1898) Wetenschappelijke Vergadering. Tijdschrift der Nederlandsche Dierkundige Vereniging 5(4), VII-IX. Weidner H (1959) Die Entomologischen Sammlungen des Zoologischen Staatsinstituts und Zoologischen Museums Hamburg. I. Teil, Pararthropoda und Chelicerata. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 57: 89-142. Wheeler WM (1898) A new Peripatus from Mexico. Journal of Morphology 15: 1-8. doi: 10.1002/jmor.1050150102 Willey A (1898a) The anatomy and development of Peripatus novae-britanniae. The University Press, Cambridge, England, 52 pp. - Willey A (1898b) On Peripatus novae-britanniae, sp. n. Annals and Magazine of Natural History [Series 7] 1: 286–287. - Wood-Mason J (1879) Morphological notes bearing on the origin of insects. Transactions of the Entomological Society of London [1879] 2: 145–167. + Willey A (1898b) On Peripatus novae-britanniae, sp. n. Annals and Magazine of Natural History [Series 7] 1: 286–287. + Wood-Mason J (1879)Morphological notes bearing on the origin of insects. Transactions of the Entomological Society of London [1879] 2: 145–167. Zilch A (1954a) Onychophoren aus El Salvador. Senckenbergiana Biologica 35: 147-150. Zilch A (1954b) Ein neuer Oroperipatus aus Peru. Senckenbergiana Biologica 35: 151-154. From 8f8ffec6ffba4d170f6e653a0e31f57b46156e72 Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 24 Mar 2014 08:05:14 +0000 Subject: [PATCH 15/18] Added 10.3897/zookeys.345.5840 --- 10.3897:zookeys.345.5840.xml | 967 +++++++++++++++++++++++++++++++++++ 1 file changed, 967 insertions(+) create mode 100644 10.3897:zookeys.345.5840.xml diff --git a/10.3897:zookeys.345.5840.xml b/10.3897:zookeys.345.5840.xml new file mode 100644 index 0000000..86d518b --- /dev/null +++ b/10.3897:zookeys.345.5840.xml @@ -0,0 +1,967 @@ +
+ + + ZooKeys + + ZooKeys + ZooKeys + + 1313-2989 + 1313-2970 + + Pensoft Publishers + + + + 10.3897/zookeys.345.5840 + + Taxonomy, distribution, and natural history of flying foxes (Chiroptera, Pteropodidae) in the Mortlock Islands and Chuuk State, Caroline Islands + + + + + Buden + Donald W. + + 1 + + + + Helgen + Kristofer M. + + 2 + + + + Wiles + Gary J. + + 3 + + + + Division of Natural Sciences and Mathematics, College of Micronesia–FSM, P.O. Box 159, Kolonia, Pohnpei 96941, Federated States of Micronesia + + + Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013-7012, USA + + + Washington Department of Fish and Wildlife, 600 Capitol Way North, Olympia, Washington 98501-1091, USA + + + +

Corresponding author: Donald W. Buden (don_buden@comfsm.fm)

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Academic editor: W. Bogdanowicz

+ + + + 2013 + + + 29 + 10 + 2013 + + 345 + 97 + 135 + + + 20 + 6 + 2013 + + + 17 + 10 + 2013 + + + + Donald W. Buden, Kristofer M. Helgen, Gary J. Wiles + + This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. + + + + +

The taxonomy, biology, and population status of flying foxes (Pteropus spp.) remain little investigated in the Caroline Islands, Micronesia, where multiple endemic taxa occur. Our study evaluated the taxonomic relationships between the flying foxes of the Mortlock Islands (a subgroup of the Carolines) and two closely related taxa from elsewhere in the region, and involved the first ever field study of the Mortlock population. Through a review of historical literature, the name Pteropus pelagicus Kittlitz, 1836 is resurrected to replace the prevailing but younger name Pteropus phaeocephalus Thomas, 1882 for the flying fox of the Mortlocks. On the basis of cranial and external morphological comparisons, Pteropus pelagicus is united taxonomically with Pteropus insularis “Hombron and Jacquinot, 1842” (with authority herein emended to Jacquinot and Pucheran 1853), and the two formerly monotypic species are now treated as subspecies — Pteropus pelagicus pelagicus in the Mortlocks, and Pteropus phaeocephalus insularis on the islands of Chuuk Lagoon and Namonuito Atoll. The closest relative of Pteropus pelagicus is Pteropus tokudae Tate, 1934, of Guam, which is best regarded as a distinct species. Pteropus pelagicus pelagicus is the only known resident bat in the Mortlock Islands, a chain of more than 100 atoll islands with a total land area of <12 km2. Based on field observations in 2004, we estimated a population size of 925–1,200 bats, most of which occurred on Satawan and Lukunor Atolls, the two largest and southernmost atolls in the chain. Bats were absent on Nama Island and possibly extirpated from Losap Atoll in the northern Mortlocks. Resident Mortlockese indicated bats were more common in the past, but that the population generally has remained stable in recent years. Most Pteropus phaeocephalus pelagicus roosted alone or in groups of 5–10 bats; a roost of 27 was the largest noted. Diet is comprised of at least eight plant species, with breadfruit (Artocarpus spp.) being a preferred food. Records of females with young (April, July) and pregnant females (July) suggest an extended breeding season. Pteropus pelagicus pelagicus appears most threatened by the prospect of sea level rise associated with global climate change, which has the potential to submerge or reduce the size of atolls in the Mortlocks. Occasional severe typhoons probably temporarily reduce populations on heavily damaged atolls, but hunting and ongoing habitat loss are not current problems for the subspecies.

+ + + + Pteropus phaeocephalus + P. pelagicus + P. insularis + P. tokudae + Mortlock Islands + Chuuk + Micronesia + atoll + taxonomy + distribution + status + natural history + climate change + sea level rise + + + + + + Introduction +

Many islands of the west-central Pacific Ocean remain poorly known biologically, particularly the numerous, small, low-lying, coralline atolls and atoll-like islands of Micronesia. Their inaccessibility and relatively depauperate biotas (compared with those of larger and higher islands) have contributed to a paucity of visiting biologists. However, an understanding of the biogeography and biodiversity of Oceania remains incomplete without knowledge of the species that inhabit these miniscule lands.

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The genus Pteropus is comprised of about 65 species of flying foxes, making it by far the largest genus in the family Pteropodidae (Simmons 2005; Helgen et al. 2009). Primarily island-dwelling, the genus is widespread through the Indo-Pacific region westward to the islands off eastern Africa. Much taxonomic work is still needed in the genus. Additionally, many species of Pteropus remain poorly known in terms of their population status, biology, and specific conservation needs. In particular, few studies of atoll-dwelling populations have been conducted (Dolbeer et al. 1988, Wiles et al. 1991, Holmes et al. 1994). The need for such studies is especially pressing in Pacific archipelagos, where larger bats have suffered considerable declines, extirpations, and extinctions over the past 200 years (Flannery 1995; Wiles et al. 1997; Helgen 2005; Helgen et al. 2009; Wiles and Brooke 2009, Nakamoto et al. 2012).

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Although Pteropus phaeocephalus Thomas, 1882 was originally described as a distinct species endemic to the Mortlock Islands, a group of atolls in central Micronesia, there has long been recognition of the similarity between this form and Pteropus insularis, which is restricted to the neighboring main islands and barrier reef islands of Chuuk Lagoon and Namonuito Atoll located 171 km to the northwest (Oustalet 1895, Andersen 1912, Rainey and Pierson 1992, Kepler 1994, Flannery 1995). This has resulted in suggestions that Pteropus phaeocephalus may be better regarded as a subspecies or synonym of Pteropus insularis (Oustalet 1895; Koopman 1993; K. Helgen, in Simmons 2005). Information on the distribution, relative abundance, and ecology of Pteropus phaeocephalus is almost nonexistent, and the lack of comparative material from these islands, hitherto limited to the holotype, has impeded taxonomic appraisal. Specimens obtained from Namoluk and Satawan Atolls during this study provide material for new comparisons and a taxonomic reappraisal. Our field observations together with information provided by Mortlockese islanders furnish new data on the distribution, abundance, and biology of flying foxes in the Mortlocks. In light of the many threats to biodiversity on Pacific islands (Helgen et al. 2009, Wiles and Brooke 2009, Woinarski 2010), we also discuss conservation concerns relating to these bats.

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The Mortlock Islands (07°00'N, 152°35'E to 05°17'N, 153°39'E) are a part of Chuuk (formerly Truk) State, one of four states comprising the Federated States of Micronesia (FSM), the others being Yap, Pohnpei, and Kosrae. The FSM together with the Republic of Belau (Palau) make up the Caroline Islands in the tropical western Pacific Ocean. The Mortlocks are a chain of five atolls and one low, coral island spanning 224 km (Figure 1). Land area totals 11.9 km2, distributed among more than 100 islands; Ta, Satawan Atoll, is the largest island (Table 1). Maximum elevations are only 3–5 m asl. Over the years, a confusing array of alternative names and spellings has been proposed for different islands and island groups in the Mortlocks and Chuuk Lagoon (Baker 1951, Bryan 1971, Berg 1993). We use the names Northern Mortlocks for Nama Island and Losap Atoll, Central Mortlocks for Namoluk Atoll, and Southern Mortlocks for Ettal, Lukunor, and Satawan Atolls.

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Location map for the Mortlock Islands and Chuuk Lagoon, Micronesia. Inset A location of islands in the west-central Pacific Ocean, G = Guam, K = Kosrae, NG = New Guinea, Pa = Palau, Po = Pohnpei, Y = Yap; inset B southern end of Satawan Atoll, solid circles indicate beach sites where interisland movement of flying foxes was assessed (see Table 5) and the open star indicates the airport station count site on Ta Island.

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Statistical data for the Mortlock Islands, Chuuk State, Federated States of Micronesia.

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Island groupLand area (km2)aNumber of islandsLargest island (km2)aNumber of inhabited islandsNumber of residentsbDistance to next atoll (km)cObservation daysd
Northern Mortlocks
Nama Island0.751Nama (0.75)1995147
Losap Atoll1.0310Lewel (0.56)28751101
Central Mortlocks
Namoluk Atoll0.835Namoluk (0.31)14075311
Southern Mortlocks
Ettal Atoll1.8920eEttal (0.97)126773
Satawan Atoll4.5965fTa (1.55)42,935870
Lukunor Atoll2.8218Lekiniochg (1.28)21,432-26
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a From Bryan (1971). b Based on the 2000 national census (Division of Statistics 2002). c Measured reef to reef. d Total number of days spent on the island(s) by DWB while conducting faunal surveys for flying foxes, birds, lizards, butterflies, and dragonflies. e Number of islands counted by DWB while walking on the reef flat, but Bryan (1971) recorded 18. f Based on information given DWB by residents of Satawan Atoll, but exact number uncertain. Bryan (1971) indicated “approximately” 49 islands in the summary section for “Truk District,” but mentioned at least 80 named and unnamed islands in the atoll and described one area in the northeast part of the atoll as having “numerous small cays on edge of reef” without naming or numbering them. g Formerly Lukunor Island, and known also as Likinioch, Lukinoch, and Lukunoch Island.

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The Mortlocks fall within the equatorial rainbelt and are wet enough to support mesophytic vegetation (Mueller-Dombois and Fosberg 1998), though some of the smaller islets that lack a fresh water lens are more xeric. Coconut (Cocos nucifera L.) forest is the predominant vegetation type, with breadfruit (Artocarpus spp.) being a codominant tree in the interior of the larger islands. Other common forest and forest edge trees include Barringtonia asiatica (L.) Kurz, Ficus spp., Guettarda speciosa L., Hernandia sonora L., Neisosperma oppositifolia (Lam.) Fosb. & Sachet, Pandanus spp., and Terminalia samoensis Rech. Forest canopy height ranges from 10 to 20 m. The forest abuts the beach or merges abruptly with a narrow zone of coastal scrub or thicket dominated by Tournefortia argentea L. f. and Scaevola taccada (Gaertn.). Large community-maintained taro patches occupy much of the interior of Nama, Namoluk, Satawan, Kuttu, Moch, Oneop, and Lekinioch Islands. More detailed accounts of island vegetation and physiognomy exist for Losap Atoll (Severance 1976, Manner and Sana 1995), Namoluk Atoll (Marshall 1975), Ettal Atoll (Nason 1970), Lukunor Atoll (Borthwick 1977, Tolerton and Rauch n. d. [1949?]), and Kuttu Island, Satawan Atoll (Reafsnyder 1984).

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All six island groups comprising the Mortlocks are inhabited, but only 1–4 islands in each group have permanent settlements (Table 1). The other islands are visited by atoll residents with varying degrees of frequency to cultivate taro or to gather coconuts, crabs, and other forest and coastal commodities. A total of 6,911 people lived in the Mortlocks in 2000, representing a density of 581 people per km2.

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Museum specimens utilized in this study are deposited in the collections of the American Museum of Natural History, New York, USA (AMNH), the Academy of Natural Sciences, Drexel University, Philadelphia, USA (ANSP), the Natural History Museum, London, UK (BMNH), Brigham Young University-Hawaii Campus, Laie, Hawaii, USA (BYUH), College of Micronesia-FSM, Kolonia, Pohnpei, Federated States of Micronesia (COM), the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (MCZ), the Muséum National d’Histoire Naturelle, Paris, France (MNHN), the National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (USNM), and the Museum für Naturkunde, Humboldt University, Berlin, Germany (ZMB).

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Measurements of specimens are in millimeters and grams. Cranial and forearm measurements were taken on museum specimens (skins, skulls, and fluid preserved specimens) using dial calipers. Cranial measurements (see Figure 2) are abbreviated (and defined) as follows: GLS (greatest length of skull; distance from posterior midpoint on occipital crest to anterior midpoint of premaxillae); PL (palate length; distance from anterior midpoint of premaxillae to posterior midpoint of bony palate); ZW (greatest width across the zygomatic arches); IOB (width of interorbital constriction); BBC (breadth of braincase, measured across the braincase at squamosal bases); MTR (length of maxillary toothrow, alveolar distance from anterior root of canine to posterior root of last molar); CC (external, alveolar distance across upper canines); M1M1 (external, alveolar distance across upper first upper molars). Head-body length (snout to rump) and ear length were measured with a millimeter ruler on fresh material or recorded from museum specimen labels. Body mass was obtained from fresh specimens using Pesola scales or recorded from specimen labels. Immatures are defined as neonates to young weighing ≤100 g. Subadults are defined as adult-sized or nearly adult-sized young with incompletely fused skull sutures/synchondroses (cf. Helgen 2004) and some indication of cartilaginous epiphyseal swellings in the bones of the wing. Because sexual dimorphism is negligible in these bats, measurements for both sexes are pooled in our comparisons. Statistical methods included a two-sample t-test processed by Excel 2000 data analysis.

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Cranial measurements employed in this study. See Methods for abbreviations.

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Standard descriptive statistics (mean, standard deviation, and observed range) were calculated for the samples of populations and species listed in the tables. Only cranial and dental measurements were incorporated in the multivariate analyses. Principal component analyses were computed using the combination of cranial and dental measurements indicated in tables and in the text. All measurement values were transformed to natural logarithms prior to multivariate analysis. Principal components were extracted from a covariance matrix. Variables for multivariate analyses were selected judiciously to maximize sample sizes for comparison by allowing for inclusion of partially broken skulls in some cases. The software program Statistica 8.0 (Statsoft Inc., Tulsa, Oklahoma, USA) was used for multivariate analysis.

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Field work for this study began as part of a biological survey of terrestrial vertebrates and selected insect groups on Satawan Atoll, but was expanded in 2004 to include broader population assessments of flying foxes and other wildlife in the Mortlocks. Field surveys were conducted on Satawan Atoll during 17–26 December 2002, 7 July–1 August 2003, 30 March–9 April 2004, 22 June–6 July 2004, and 1–5 August 2004. The five other groups of islands were visited in 2004: Nama Island, 7–14 July; Losap Atoll, 10 July; Namoluk Atoll, 19–29 July; Ettal Atoll, 30 July–1 August; and Lukunor Atoll (Lekinioch Island only), 2–3 August. In 2012, incidental observations were made at Lukunor Atoll, 20 June–15 July, and Satawan Atoll, 15–17 July. All field work was conducted by DWB.

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Information on the abundance and biology of bats came from a combination of sources. Station counts of flying bats were conducted on Satawan Atoll at two types of locations providing relatively unobstructed views of the sky: the Ta airport and six beach sites where interisland movement of bats was assessed (Figure 1). These counts were made by a single person at sunrise or sunset, lasted 25–95 minutes, and were made once or twice per time period at each station except for multiple counts at the Ta airport.

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Flying foxes (and other wildlife) were also counted during walks conducted from mid-morning to mid- to late afternoon. These were made along pre-existing trails or by following a compass bearing through the center of less frequently visited islands, usually along the long axis of an island, but occasionally at right angles to it. Each route was surveyed only once. Care was taken to avoid double counting bats flushed by the observer. An undetermined, but probably substantial, proportion of roosting bats was hidden from view by the relatively dense forest canopy. Roosting flying foxes were viewed with 10x binoculars to observe behavior and search for the presence of young. Other information on bats was obtained from incidental observations and atoll residents. We did not search for feeding evidence of bats (i.e., discarded fruit, chewed pellets, fecal splats). An estimate of the number of flying foxes on each atoll was made based on: 1) overall numbers of bats seen, 2) percentage of each atoll covered by the observer, 3) the quality and amount of forest on an atoll, and 4) information provided by island residents, especially for Namoluk, Ettal, and Lukunor Atolls.

+ + + + Results + + Taxonomy. <italic><tp:taxon-name>Pteropus pelagicus</tp:taxon-name></italic> Kittlitz, 1836. Taxonomic history +

The Russian research vessel Senyavin, under command of Friedrich [Feodor] Lütke, and with F. H. von Kittlitz as one of the ship’s naturalists, was in the eastern Caroline Islands early in 1828 (Nozikov 1946). After leaving Ualan (= Kosrae), the Senyavin spent approximately the first two weeks of February in the southern Mortlocks (Lütke 1835). Although the ship traveled along the coasts of all three southern atolls, the only anchorage and the base of scientific operations was the harbor at Lukunor Atoll (Lütke 1835). Remarking on the fauna of Lukunor, Kittlitz (1836) stated “Nous prîme, comme à Ualan, deux mammifères dont l’un est un Pteropus (Pteropus pelagicus m.) qui sans aucun doute diffère de celui de cette île, tant par sa taille qui est beaucoup moindre, que par une tache blanche de forme circulaire qu’il a sur l’abdomen [We found, as on Kosrae, two mammals, one of which is a Pteropus (Pteropus pelagicus mihi) that without any doubt differs from the one of that island by its much smaller size and by a round white patch on its abdomen].” Although Kittlitz’s description is brief and unaccompanied by illustration, clearly he was referring to the same species that Thomas (1882: 756) named Pteropus phaeocephalus. The zoological specimens that Kittlitz obtained during this expedition were deposited in the Russian Academy of Sciences in St. Petersburg (Baker 1951, Hume 2001), but the only flying foxes in the St. Petersburg collection today are two Pteropus pselaphon Say from the Bonin Islands, Japan (V. Loskot, pers. comm.). We thus regard Pteropus pelagicus as the earliest name for the flying fox currently known as Pteropus phaeocephalus, and advocate its usage (see Taxonomic Conclusions, below).

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Thomas (1882) described Pteropus phaeocephalus from the Mortlock Islands based on “a gravid female in alcohol” collected by Dr. [John = Johann = Jan] Kubary for the Godeffroy Museum, Hamburg, Germany, and subsequently transferred to the Natural History Museum, London (BMNH 82.10.27.4). Andersen (1912: 299) considered the holotype as “perfectly similar to Pteropus insularis [from Chuuk Lagoon islands] in skull, dentition, palate-ridges, and all external characters, except much paler colour of the fur, particularly on the back.” He (Andersen 1912: 299) concluded “the evidence is in favour of the assumption that Pteropus phaeocephalus (Mortlock group) is specifically distinct from Pteropus insularis (Ruck [= Chuuk] group).” This opinion has prevailed in the literature until relatively recently. Koopman (1993) remarked that Pteropus phaeocephalus is probably better recognized as a subspecies of Pteropus insularis, and Simmons (2005) concurred in referring to the Mortlock population as Pteropus insularis phaeocephalus.

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The holotype of Pteropus phaeocephalus is labeled as collected by Kubary in the Mortlock Islands, but the date and the name of the island are unstated. Kubary traveled widely in the Pacific while employed as a naturalist and ethnographer by the Godeffroy Museum (Spoehr 1963). He visited the Mortlock Islands apparently on two occasions. Hezel (1979: 33, revised online as of 2002 and unpaged) stated that the brig Iserbrook put in at Ta, Satawan Atoll on 16 February 1873 and “returned in July or August with John Kubary aboard from Palau.” Referring to the same voyage, Paszkowski (1971: 48) stated that “in May, 1873, Kubary sailed on the Iserbrook [from Palau] and visited the islands of Ngulu, Ulithi, Woleai, Nukuoro, and Mortlock…[and] landed at Ponape [= Pohnpei] in August, 1873.” Little is known of Kubary’s activities in the Mortlocks during this visit, which may have been very brief, and Kubary did not allude to it in his ethnography of the Mortlocks (Kubary 1880). Also, any specimens that may have been collected at this time may have been among those lost in a subsequent shipwreck at Jaluit, in the Marshall Islands, in September 1874, when only a small part of Kubary’s collections of the previous five years were saved (Paszkowski 1971, 1987).

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Kubary returned to the Mortlocks in 1877. In a separate and anonymously written introduction to the ethnography that Kubary apparently sent to L. Friederichsen, editor of Mittheilungen der Geographischen Gesellschaft in Hamburg, the author (L. Friederichsen?) or authors (in Kubary 1880) stated “Kubary verweilte auf den Mortlock-Inseln während der Monate März bis Ende Mai 1877 und zwar speciell auf der Inseln Tä, Uoytä und Aliar [Kubary visited the Mortlock Islands, particularly the islands Ta, Weito, and Aliar during the months of March through the end of May 1877].” Westwood (1905: 89, 112) saw Kubary also on Satawan Island (immediately to the east of Ta) during his visit in 1877. In light of this information, the holotype of Pteropus phaeocephalus probably was collected on Satawan Atoll sometime during March–May 1877 and very likely on Ta Island, where flying foxes are now common and where Kubary seems to have spent most of his time during his three months in the Mortlocks (Nason 1970: 363).

+

Matschie (1899) discussed a specimen he referred to as Pteropus phaeocephalus from Pohnpei in the Caroline Islands, obtained by Otto Finsch (ZMB A4065), but this is actually a specimen of a Pteropus allied to Pteropus mariannus (Andersen 1912), previously discussed as a specimen of Pteropus ualanus, otherwise known only from Kosrae (Wiles et al. 2008) but in fact most closely resembling Pteropus pelewensis of Palau (confirmed by KMH). The source of Finsch’s specimen was possibly mislabeled or the specimen may represent a population of Pteropus pelewensis (or a closely related taxon) that has become extinct on Pohnpei, where only Pteropus molossinus occurs today (Flannery 1995).

+ + + Morphological comparisons. External comparisons +

Among the 11 specimens from Satawan Atoll, the mantle ranges from creamy white to deep buff or tawny in adults and subadults, and is grayish brown (without yellowish or reddish highlights) in immatures. It averages darker (more tawny, less pale buff) in the six adults from Namoluk Atoll. In all Mortlock specimens, the mantle is bisected by a narrow to broad area of darker (brown) pigmentation, which, in some cases, gives the appearance of having pale shoulder patches. In specimens from both Namoluk and Satawan, the back and rump are dark brown (occasionally tinged with reddish brown in Satawan specimens), with scattered pale-tipped hairs; Namoluk specimens average slightly darker than those from Satawan. In both the Namoluk and Satawan material, the face is dark brown to nearly black and with varying amounts of grizzling. The top of the head is brown or grayish brown. The venter is usually light brown to pale reddish brown anteriorly (on throat and chest) and dark brown posteriorly; subadults are more uniformly light brown. A large prominent white or pale yellowish white patch, 20–40 mm wide, occurs midventrally in most specimens (Figure 3C); it is occasionally slightly reduced or nearly obsolete.

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(A) Scanned image of Plate LIV from original description of Pteropus phaeocephalus Thomas 1882, which is presumably based on the holotype (B) dorsal view and (C) ventral view of Pteropus pelagicus pelagicus from Satawan Atoll (D) ventral view and (E) dorsal view of Pteropus phaeocephalus insularis from Weno, Chuuk Lagoon.

+ + +

Mortlock Islands specimens generally match the color descriptions of the holotype of phaeocephalus as described by Thomas (1882) and Andersen (1912), although the pale-tipped hairs on the dorsum described as “nearly white” by Thomas (1882) and “golden cream buff” by Andersen (1912) are much sparser in the new material and visible only upon close inspection. Mortlock specimens, especially some of the paler and brighter individuals from Satawan, closely resemble the illustration (Plate LIV) in Thomas (1882) (see Figure 3A). DWB observed an unusually pale individual through binoculars from a distance of 30–40 m on Toimon Island, Namoluk Atoll, on 27 July 2004. The venter and sides were entirely creamy white. Only the top of the head, neck, a small part of the mantle middorsally, and most of the back and rump were dark brown.

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Compared with a sample of five specimens of Pteropus insularis, all presumably from Chuuk Lagoon (four from Weno, one [MCZ 7023] of uncertain origin), the Mortlock Islands bats tend to show greater contrast between the paler coloration of the mantle and the darker pigmentation on the back and rump. The midventral pale patch is consistently larger in the Mortlock specimens that we examined, although Andersen (1912: 296) stated “the size of this bright pectoral patch varies greatly individually [in Pteropus insularis from Chuuk Lagoon], but in none is it wanting….” The cotype of Pteropus insularis from Chuuk Lagoon illustrated in Hombron and Jacquinot (1844: plate v) shows a prominent mid-ventral patch as large as any seen in specimens from the Mortlocks (George Glazer Gallery web page at www.georgeglazer.com/archives/prints/animals/durvillebat.html ). Hair length was measured only on the mantle and averaged slightly shorter in Chuuk Lagoon specimens than in those from the Mortlocks—14.7 mm long in three specimens from Weno, Chuuk Lagoon, and 15.0 and 16.0 mm in four from Namoluk and nine from Satawan, respectively. In body size, bats from Chuuk Lagoon are similar to those from the Mortlocks. One skin and skull (AMNH 249954) dated 26 May 1987 and 13 other skulls (AMNH 249955–67) dated 3 July 1986 were shipped from Chuuk State and intercepted by customs officials in Guam; the specimens were prepared by G. Wiles. Whether the AMNH series consists entirely of specimens from Chuuk Lagoon, which is likely, or whether it includes material from the Mortlocks or other Chuuk outliers, is unknown. During the 1980s, bats commercially hunted in the Mortlocks were transshipped to Weno for transfer to Guam and the Commonwealth of the Northern Mariana Islands together (but apparently unsegregated) with bats collected on islands in Chuuk Lagoon. In external measurements, the AMNH specimens are similar to samples from both Weno and those from the Mortlocks (Table 2). Chromatically, the lone AMNH skin more closely resembles those from Satawan in the Mortlocks than it does the small sample from Weno.

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Mensural data for samples of flying foxes from Chuuk Lagoon islands and the Mortlock Islands; data sets include range, n in parentheses, and mean ± SD; all measurements are in millimeters or grams.

+ + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
Mortlock Islands
CharacterSex“Truk”aChuuk Lagoon islandsNamoluk AtollSatawan Atoll
Head-body length♂____168.0–186.0 (2) 177.0 ± 12.73158.0 (1)150.0–170.0 (2) 160.0 ± 14.0
♀165.0 (1)131.0 (1)155.0–170.0 (5) 163.0 ± 6.08155.0–170.0 (4) 161.3 ± 6.29
Forearm length♂____101.0 (1)103.9 (1)102.0–108.7 (2) 105.4 ± 4.74
♀104.8 (1)____102.3–107.3 (5) 104.6 ± 2.18103.2–108.0 (5) 105.3 ± 1.78
Ear length♂20.0–22.0 (2) 21.0 ± 1.4123.0 (1)24.0 (1)
♀________23.0–24.0 (3) 23.7 ± 0.5822.0–23.0 (2) 22.5 ± 4.74
Body mass♂____148.0–200.0 (2) 174.0 ± 36.76175.0 (1)142.0–203.0 (2) 172.5 ± 43.1
♀170.0 (1)150.0 (1)190.0–225.0 (5) 211.0 ± 13.87153.0–187.0 (4) 171.8 ± 12.28
Greatest skull length (GLS)♂43.9–46.1 (5) 44.8 ± 0.18 44.6 (1)45.0 (1)47.2 (1)
♀44.0–45.4 (5) 44.7 ± 0.5344.1 (1)45.3–46.1 (2) 45.7 ± 0.5745.0–46.6 (4) 45.7 ± 0.59
Palate length (PL)♂22.6–24.0 (5) 23.0 ± 0.5922.2 (1)23.3 (1)22.0–24.4 (2) 23.2 ± 1.70
♀22.0–24.0 (6) 22.8 ± 0.7522.9 (1)23.0–24.0 (2) 23.5 ± 0.7123.2–24.1 (3) 23.9 ± 0.59
Maxillary toothrow (MTR)♂14.8–15.2 (5) 15.1 ± 0.1715.1–15.2 (2) 15.2 ± 0.7015.5 (1)15.2–15.8 (2) 15.6 ± 0.50
♀15.0–15.5 (6) 15.2 ± 0.1914.8 (1)14.8–15.5 (3) 15.2 ± 0.3615.0–15.5 (4) 15.2 ± 0.24
Breadth of braincase (BBC)♂17.2–17.6 (5) 17.4 ± 0.1517.3 (2)18.0 (1)17.6–18.2 (2) 17.9 ± 0.42
♀17.0–17.8 (6) 17.5 ± 0.3317.0 (1)17.9–18.1 (2) 18.0 ± 0.1417.5–18.1 (4) 17.9 ± 0.26
Breadth across canines (CC)♂10.0–11.0 (5) 10.4 ± 0.4410.8–11.1 (2) 11.0 ± 0.2111.1 (1)10.7–11.9 (2) 11.3 ± 0.85
♀9.6–10.8 (6) 10.4 ± 0.419.9 (1)11.0–11.4 (3) 11.2 ± 0.2110.9–11.1 (4) 11.0 ± 0.10
Breadth across M1 (M1M1)♂12.0–12.5 (4) 12.3 ± 0.2212.2 (2)12.0 (1)12.3–12.9 (2) 12.6 ± 0.42
♀12.1–12.8 (5) 12.3 ± 0.2912.0 (1)11.9–12.3 (3) 12.1 ± 0.2112.3–12.8 (3) 12.6 ± 0.26)
Interorbital breadth (IOB)♂7.0–7.4 (4) 7.2 ± 0.177.5 (1)7.5 (1)7.1–7.6 (2) 7.4 ± 0.35
♀7.0–7.4 (6) 7.2 ± 0.147.2 (1)7.7–8.0 (3) 7.8 ± 0.157.7–7.8 (3) 7.8 ± 0.06
Zygomatic width (ZW)♂23.0–25.4 (4) 24.1 ± 0.9524.0–27.0 (2) 25.5 ± 2.2125.6 (1)23.9–27.1 (2) 25.5 ± 2.26
♀23.6–25.4 (5) 24.5 ± 0.65____25.7–26.0 (2) 25.9 ± 0.2125.2–25.6 (3) 25.4 ± 0.20
+ + +

a Specimens confiscated on Guam, but originated from Chuuk State. Most were probably collected on islands in the Chuuk Lagoon, but some may have come from outer atolls, including the Mortlocks.

+ + + +

The type series of Pteropus laniger, originally described as Pteropus lanigera [sic]by H. Allen (1890), and now regarded as a synonym of Pteropus phaeocephalus insularis (Andersen 1912; Helgen et al. 2009) (USNM 37815, lectotype, and MCZ 7023, paralectotype), was stated by Allen (1890) to have come from Samoa, but this was later emended to the Caroline Islands (see Andersen 1912; Helgen and McFadden 2001). These specimens match well with BYUH specimens from Weno in being nearly uniformly brown with less contrasting areas of coloration, and having a reduced midventral pale patch, and probably originated from one of the main islands of Chuuk Lagoon.

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Overall pelage coloration of the two available skins of Pteropus tokudae (e.g. Figure 4; see further discussion below) is generally similar to that of Pteropus phaeocephalus insularis, perhaps averaging somewhat darker overall (Tate 1934; Flannery and Schouten 2001: 172).

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Holotype skin of Pteropus tokudae. AMNH 87117, adult male, from Guam, collected 10 Aug 1931 by W.F. Coultas. Scale bar = 25 mm.

+ + + + + + Craniometric comparisons +

Our study of essentially all available museum specimens of Pteropus from the islands of Chuuk Lagoon, Namonuito Atoll, and the Mortlocks, including the newly collected Mortlock material reported here, allowed us to examine patterns of multivariate craniometric variation in Pteropus from these archipelagos using Principal Component Analyses (PCA). We also included in these comparisons the three available skulls of Pteropus tokudae Tate, 1934, of Guam (a species that apparently became extinct in the late twentieth century: Wiles 1987a), which appears to be the closest relative of Pteropus pelagicus. Pteropus tokudae and Pteropus pelagicus share most qualitative morphological features but purportedly differ in that Pteropus tokudae is darker brown in coloration and smaller in body size (Tate 1934; Flannery 1995); Tate (1934) indicated in the original description of tokudae that, “It may well be merely a race of insularis.” We included Pteropus tokudae in these comparisons because no study has yet explored the consistency of differences between Pteropus tokudae and Pteropus pelagicus, and because, as noted above, the largest available sample of Pteropus specimens supposedly originating from Chuuk State was seized in Guam, such that we wanted to evaluate whether these specimens showed closer craniometric relationship with Pteropus phaeocephalus insularis of Chuuk Lagoon, or with Pteropus tokudae of Guam. Demonstration of the precise phylogenetic relationships of Pteropus pelagicus and Pteropus tokudae awaits a more detailed molecular phylogenetic review of the genus Pteropus than has yet been published (Giannini et al. 2008), but we regard Pteropus pelagicus and Pteropus tokudae to be closely related species that constitute a small radiation of relatively small-bodied flying-foxes from islands of the remote Central Pacific.

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The type of pelagicus can no longer be traced (see above), and the skulls of the surviving type specimens for the other relevant taxa in this group (insularis [MNHN 1996-2112, syntype of insularis, designated here as the lectotype of insularis based on being the only skull available from the type series], phaeocephalus [BMNH 82.10.27.4, holotype], laniger [USNM 37815, lectotype], and tokudae [AMNH 87117, holotype] are all somewhat broken. Consequently, we based our PCA comparisons on a relatively small number of five variables (Table 4, Figure 5) to allow for the inclusion of these various type specimens, as well as various other museum specimens (adult skulls at AMNH, ANSP, BMNH, COM, MNHN, USNM, and ZMB), many of which are also partially broken.

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Cranial measurements (mean ± SD, in millimeters, n in parentheses) for pooled samples of flying foxes from Chuuk Lagoon islands versus Mortlock Islandsa.

+ + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
CharacterChuuk LagoonbMortlock IslandstdfP
Greatest skull length44.8 ± 0.63 (12)45.9 ± 0.80 (6)-3.37180.003
Palate length22.8 ± 0.64 (13)23.5 ± 0.82 (8)-2.22190.039
Maxillary toothrow15.1 ± 0.18 (14)15.3 ± 0.32 (10)-1.93220.066
Breadth of braincase17.4 ± 0.26 (14)17.9 ± 0.24 (9)-4.88210.000
Breadth across canines10.5 ± 0.44 (14)11.1 ± 0.34 (10)-3.93220.001
Breadth across M112.4 ± 0.30 (12)12.3 ± 0.36 (9)0.21190.834
Interorbital breadth7.2 ± 0.16 (12)7.7 ± 0.26 (9)-4.83190.000
Zygomatic width24.5 ± 1.07 (11)25.6 ± 0.89 (8)-2.21170.041
+ + +

a Unpooled sample statistics are in Table 2. b Includes AMNH specimens confiscated on Guam.

+ + + + + +

Morphometric separation (first two principal components of a Principal Components Analysis) of 27 adult skulls of Pteropus pelagicus and Pteropus tokudae. These comparisons involve 5 measurements (maxillary toothrow length, breadth of braincase, external breadth of rostrum across canines, external breadth of palate across first upper molars, and least interorbital breadth). The first principal component mainly reflects distinctions in overall skull size, which increases from right to left. Specimens of Pteropus pelagicus pelagicus from the Mortlock Islandsare denoted by red symbols (including the red triangle, the holotype of phaeocephalus from “Mortlock Islands”; red circles, Satawan Atoll; and red squares, Namoluk Atoll). Specimens of Pteropus pelagicus insularis are denoted by black symbols; closed black symbols indicate samples of known geographic origin (including the closed black triangle, the holotype of insularis from “Ruck”; closed black circles, specimens labeled “Ruck”; closed black square, specimen labeled “Uala” (= Weno); and black cross, specimen from Namonuito Atoll) and open symbols indicate specimens of imprecise geographic origin (including the large open triangle, the lectotype of laniger, erroneously attributed to the “Samoa Islands” in the original description; open black circle, an unprovenanced specimen from ANSP; and open black diamonds, specimens at AMNH seized on Guam but originating from Chuuk State). Specimens of the closely related species Pteropus tokudae from Guam are denoted by pink symbols (including the pink triangle, the holotype of tokudae from Guam, and the pink squares, other specimens from Guam).

+ + +

Our craniometric comparisons (Figure 5, Table 4) indicate that skulls from the Mortlock Islands (pelagicus), from the islands of Chuuk Lagoon and Namonuito Atoll (insularis), and from Guam (tokudae), separate primarily along the first principal component (representing 71% of the variance), mainly reflecting differences in overall skull size between these insular populations/taxa.

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Factor loadings, eigenvalues, and percentage of variance explained by illustrated components (Figure 5) from Principal Components Analysis of 27 adult skulls of Pteropus pelagicus and Pteropus tokudae (see “specimens examined”). Principal components are extracted from acovariance matrix of 5 log-transformed cranialmeasurements (see Figures 2 and 5, Tables 2–3).

+ + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
VariablePC1PC2
Interorbital width-0.5870.798
Breadth of braincase-0.869-0.121
Maxillary toothrow length-0.911-0.163
Breadth across canines-0.972-0.105
Breadth across M1s-0.711-0.368
Eigenvalue0.0130.003
Percent variance71.4%17.9%
+ +

Several points are of special note. First, specimens of tokudae separate cleanly from pelagicus/insularis, chiefly on the basis of their consistently smaller size (showing no overlap with Pteropus pelagicus in most skull measurements). Pteropus tokudae has a markedly smaller skull (greatest length 41.2–42.2 mm) with a relatively shorter and narrower rostrum compared to Pteropus pelagicus (Figure 6).

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Skulls of Pteropus pelagicus (A Pteropus phaeocephalus insularis, USNM 151563, unsexed adult, from Uala [= Weno]; note last lower premolars missing in lower jaw) and Pteropus tokudae (B AMNH 87117, holotype of tokudae, adult male, from Guam). Scale bar = 25 mm.

+ + +

Second, some overlap in morphometric space characterizes samples of pelagicus (skulls generally larger)and insularis (skulls generally smaller), especially with regard to the holotype of Pteropus phaeocephalus, which associates more closely with Chuuk Lagoon and Namonuito specimens than with more recent Mortlock samples. Chuuk Lagoon and Namonuito specimens tend to average slightly smaller in most measurements than Mortlock specimens (Table 2), but sample sizes are small. Statistically significant differences (p < 0.05) were found in six of eight cranial measurements when samples from different localities were pooled and sexes combined (Tables 2–3); ranges overlap in all measurements.

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The only available skull from Namonuito Atoll (BMNH 15.1.18.1) shows closest association with specimens from Chuuk Lagoon (Figure 5), an indication that the flying fox population of Namonuito, the taxonomic position of which has not previously been analyzed and has been considered uncertain (Rainey and Pierson 1992, Flannery 1995), is best referred to Pteropus phaeocephalus insularis (but its taxonomic status should ideally be re-evaluated once additional comparative material is available). The lectotype of laniger also associates craniometrically with specimens of Pteropus phaeocephalus insularis (Figure 5), supporting the suggestion that the type series of laniger originated from Chuuk Lagoon (Andersen 1912, Helgen and McFadden 2001, Helgen et al. 2009) and that laniger can be regarded as a synonym of Pteropus phaeocephalus insularis. The majority of specimens at AMNH that were seized on Guam and said to originate from Chuuk State fall within the range of morphometric variation for specimens known to have come from Chuuk Lagoon (Figure 5), supporting the idea that they likely originated from Chuuk Lagoon, and their taxonomic identification as Pteropus phaeocephalus insularis. However, the AMNH series, along with an unprovenanced specimen from the ANSP (ANSP 6196), expand the range of variation in Pteropus phaeocephalus insularis along the second principal component (representing 18% of the variance), suggesting a slightly greater range of cranial shape variation within Pteropus phaeocephalus insularis than documented by firmly localized samples. The possibility that some of these unprovenanced AMNH and ANSP specimens might have originated from the Mortlocks, or other outlying island populations of Chuuk State, cannot be ruled out.

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Our search for museum material of Pteropus phaeocephalus pelagicus and Pteropus phaeocephalus insularis uncovered one additional very interesting and largely overlooked museum specimen. This is a broken skull of a flying fox much larger than Pteropus phaeocephalus insularis, received from Otto Finsch, probably in 1880 (when he was in the Caroline Islands during his first Pacific expedition), and labeled “Ruck” (= Chuuk). This specimen (ZMB 5697) was misidentified as Pteropus insularis by Matschie (1899), who figured the skull (Matschie 1899: Plate 5, numbers 11-12), but was correctly identified by Andersen (1912) as a member of the remote Pacific group of flying foxes allied taxonomically with Pteropus mariannus (including the named forms pelewensis, yapensis, and ualanus). This specimen, examined by KMH, most closely resembles Pteropus pelewensis of Palau and provides an indication that, like Guam (the original fauna of which included Pteropus mariannus and Pteropus tokudae), the islands of the Chuuk Lagoon may have supported both a smaller species of flying fox (Pteropus phaeocephalus insularis) and Pteropus cf. pelewensis. No other collections or reports from Chuuk State give any indication of a larger species of flying fox in the archipelago, suggesting that any larger taxon that may have inhabited these islands is now extinct. However, the possibility that it could still exist in an outlying island group in Chuuk State should be kept in mind by future fieldworkers in these regions. Similarly, and as noted above, Finsch’s overlooked nineteenth century Pacific collections in Berlin reveal that the fauna of Pohnpei also may have included both a small species, Pteropus molossinus, which survives, and a larger species, also most closely resembling Pteropus pelewensis, unknown today in Pohnpei. Alternatively, we cannot fully discount the possibility that the two anomalous specimens (one from Chuuk, the other from Pohnpei) are specimens of Pteropus pelewensis from Palau that have been mislabeled. Both were obtained by Finsch and deposited in the collections of the Berlin Museum (ZMB). Finsch visited Pohnpei in 1880 (Finsch 1881), but we found no mention of his having travelled to Chuuk or Palau. However, while on Pohnpei, he visited with Johann Kubary, who resided at Pohnpei at the time and who had collected zoological specimens extensively throughout the Caroline Islands, including in Chuuk and Palau. Finsch (1880) published many new locality records of birds from Chuuk State based on Kubary’s specimens that he (Finsch) examined during this visit. Possibly the enigmatic specimen of flying fox from “Ruck” originated from Kubary’s collections. The likelihood of mislabeling in the cases of these two specimens is further enhanced by the tendency of the curators at the Berlin Museum at that time to remove collectors’ labels when attaching uniformly standardized museum labels (Frahnert and Buden 2010).

+ + + Taxonomic conclusions +

The name Pteropus pelagicus Kittlitz, 1836 is a senior synonym of the younger but prevailing name Pteropus phaeocephalus Thomas, 1882. Kittlitz’s name is listed in Sherborn’s (1929) Index Animalium (Part XIX: 4818), which Gardner and Hayssen (2004) included in a list of important nomenclators in mammalogy. It is also used in a similar context in Kramer’s (1935) report on the results of the German 1908–1910 South Seas Expedition. Inasmuch as the name Pteropus pelagicus appears in at least these two post-1899 publications, it is not a “forgotten name” (nomen oblitum) sensu ICZN (1999: Article 23.9 and glossary), and we find no compelling reason to maintain use of the younger name under special provisions of the Code (ICZN 1999: Article 23.9.3). The species known largely by its junior synonym, Pteropus phaeocephalus, has a very limited distribution, has until now been known by a single museum specimen, and has been rarely discussed in the literature. In a strict application of the Principle of Priority, the name Pteropus pelagicus Kittlitz, with type locality Lukunor Atoll in the Mortlock Islands, is resurrected here to replace the younger synonym, Pteropus phaeocephalus Thomas.

+

In view of the very close similarity in body size and cranial features and measurements between specimens from Chuuk Lagoon islands (Pteropus insularis) and those from the Mortlocks Islands (Pteropus pelagicus), we include Pteropus insularis Jacquinot and Pucheran, 1853 (herein emended from Hombron and Jacquinot, 1842 – see remarks on authorship, this account) in Pteropus pelagicus and treat the two former monotypic species as subspecies – Pteropus phaeocephalus pelagicus Kittlitz, 1836 in the Mortlock Islands (synonym phaeocephalus Thomas, 1882) and Pteropus phaeocephalus insularis Jacquinot & Pucheran, 1853 on Chuuk Lagoon islands and Namonuito Atoll (synonym laniger Allen, 1890). Nomenclatorial issues notwithstanding, this arrangement follows in principle that of Simmons (2005), who referred to the Mortlock population as Pteropus insularis phaeocephalus. The nominate subspecies (Pteropus phaeocephalus pelagicus) is distinguished from Pteropus phaeocephalus insularis chiefly, and on average, by its brighter coloration, more contrastingly colored (paler) mantle, larger white or pale buff midventral patch, and larger cranial size. The euphonious English vernacular name “Pelagic flying fox” would logically stem from the binomen, but we suggest the name “Chuuk flying fox” for this species following a tendency to use geographic-based names for the pteropodids endemic to the Caroline Islands (e.g. Flannery 1995, Wilson and Cole 2000, Simmons 2005, Wiles 2005).

+ + + Remarks on the authority of <italic><tp:taxon-name>Pteropus insularis</tp:taxon-name></italic> +

“Hombron and Jacquinot, 1842” has long been the recognized authority for the name Pteropus insularis (e.g. Andersen 1912, Wilson and Reeder 1993, Flannery 1995), but a recent review of the early literature indicates this to be an error perpetuated over the years. The name was proposed in the results of a 19th century French expedition to Antarctica and Oceania known in abbreviated form as “Voyage au Pole Sud,” or in a longer and more complete form as “Voyage au pôle sud et dans l’Océanie sur les corvettes l’Astrolabe et la Zélée exécuté par ordre du roi pendant les années 1837–1838–1839–1840 sous le commandement de M. Dumont-d’Urville.” This work was published in seven parts (Histoire du Voyage, Zoologie, Botanique, Anthropologie, Géologie, Physique, and Hydrographie) in 23 volumes with six atlases. Many authors and editors contributed to the work and publication dates range from 1842–1854 for the different components. Pteropus insularis was described in Part II (Zoologie), volume 3, which was edited by Hombron and Jacquinot and divided into 3 sections under different authors. The first section, Mammifères et Oiseaux by H. Jacquinot and Pucheran (pp. 1–166), contains the description of Pteropus insularis on page 24. The two other sections are Reptiles et Poissons by H. Jacquinot and Guichenot (pp. 1–56) followed by Crustacés by H. Jacquinot and Lucas (pp. 1–107). Clark and Crosnier (2000) remark that the earliest verifiable date for volume 3 is 1854 (although 1853 appears on the title page), but Holthuis (2002: 419) pointed out that “as there is no evidence to prove that the volume was published after 1853…the date on the title page [see Holthuis 2002: figure 7] has to be accepted as correct.”

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The description of Pteropus insularis by Jacquinot and Pucheran (1853) includes a reference to plate v, which was published in a separate Atlas d’Histoire Naturelle, Zoologie, edited by Hombron and Jacquinot. The plate is a composite of dorsal and ventral views of the body and dorsal, ventral, lateral, and frontal views of the skull of one of the two cotypes (skin 53 A, skull A. 6770) of Pteropus insularis in the Paris Museum (Andersen 1912). The plate is undated but the title page of the atlas bears the range of dates 1842–1853. However, a detailed study of the atlas by Clark and Crosnier (2000) revealed that the 40 plates were distributed in 28 livraisons, and that plate v was included in the 11th livraison, the earliest verifiable date of which is 1844, according to British Library records. But the date of the plate is seemingly irrelevant as it identifies the species with a French vernacular name, Rousette Insulaire, not a scientific binomen. On the other hand, Jacquinot and Pucheran (1853: 24) give the name as “Rousette insulaire.—Pteropus insularis Homb. et Jacq.” Although Jacquinot and Pucheran (1853: 25) remarked “La Rousette insulaire a été trouvée par MM. Hombron et Jacquinot dans l’île d’Hogoleu [The island flying fox was found by MM. Hombron and Jacquinot in Chuuk.]”, and attributed authorship to them, there is no indication that Hombron contributed directly to the description. The authority of Pteropus insularis should revert to Jacquinot and Pucheran (1853). It is also noteworthy that Sherborn’s Index Animalium, which lists original sources of species names up to 1850, does not include Pteropus insularis.

+ + + Distribution and status of <italic><tp:taxon-name>Pteropus phaeocephalus pelagicus</tp:taxon-name></italic>. Northern Mortlocks +

Records of flying foxes from the northern Mortlocks are scanty, lacking in substantive detail, and confined to Losap Atoll; no records exist for Nama Island. Severance (1976) remarked that bats were present on Losap during his anthropological investigations in the late 1960s and early 1970s, but the information may have been based on hearsay from local residents as he did not later recall definitely seeing bats on the atoll (C. Severance, pers. comm.). Kachusy Silander, whom Severance first met on Pis Island, Losap Atoll, and who lived there until the early 1970s, stated in a telephone conversation (fide C. Severance, pers. comm.) that he never saw bats on the atoll, but recalled that the traditional chief at that time claimed that flying foxes lived on his parcel of land at the southeastern end of Pis. In 2004, DWB spoke with two former Losap Island residents who had relocated to Pohnpei. Tasiro Leg (pers. comm.) recalled occasionally seeing bats and throwing stones at them as a boy during the 1980s. Scaima Pamar (pers. comm.), age 78 in 2004, recalled seeing his mother and grandmother preparing bats to eat on Pis during the time of the Japanese administration in the 1930s or early 1940s. DWB also received a report from another former Losap resident, age 76 in 2004, who remembered a large group of bats arriving together at Losap shortly after World War II (fide Tridell Elitok, pers. comm.). They resided in a no longer present stand of mangroves and were caught and eaten by the islanders because of the scarcity of food at the time. H. Manner (pers. comm.), who spent one week collecting plants on Losap Island and adjacent islets in July 1988 (Manner and Sana 1995), did not remember seeing flying foxes then.

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In the late 1980s, government officials on Weno, Chuuk Lagoon, reported bats present on Losap Atoll and said that a Losap islander was among several Mortlockese shipping bats to Weno for export to the Mariana Islands (Wiles 1992a), but whether any of the bats originated from Losap Atoll is unknown. Joakim Peter (pers. comm., College of Micronesia, Weno) informed DWB that the exporter from Losap resided on Weno at the time and worked mainly for his brother-in-law, who was from Ettal and also resided on Weno. J. Peter (pers. comm.) went on to say that the exporter from Losap may have been more involved with transportation and management operations than in collecting bats. Attempts to contact this exporter in 2004 were unsuccessful. Rainey and Pierson (1992), citing M. Henry and C. Glover in Chuuk Lagoon as sources, also reported bats from Losap but gave no additional details.

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DWB saw no bats on Losap Atoll during a one-day visit that included 2–3 hrs each on Pis, Losap, and Lewel Islands in July 2004. Lewel, the largest island in the atoll, has extensive breadfruit-coconut forest typical of bat habitat elsewhere in the Mortlocks. It is used as a garden island by the people of Losap Island, but has no permanent settlement or resident families. He walked the length of Lewel through the center and returned via a shore route without seeing bats. None of the several Losap and Pis residents interviewed during his visit knew of bats occurring on the atoll.

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DWB spent one week on nearby Nama Island without seeing any bats or encountering any resident islanders who had seen them. Mitaro Chosa (pers. comm.), deputy mayor of the island and age 58 in 2004, was firm in his conviction that bats had not occurred on Nama during his lifetime.

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These few, and in some cases questionable, records together with the lack of sightings during this study suggest that flying foxes are absent from Nama Island, and are either recently extirpated or possibly still present in such small numbers as to be unknown to many islanders on Losap Atoll.

+ + + + Central Mortlocks +

Girschner (1912) recorded flying foxes on Namoluk Atoll in about 1910, but gave no indication of their abundance. Bats were present on all of the atoll’s islands during the late 1960s and early 1970s and reportedly numbered “in the hundreds” (Marshall 1972: 17; 1975). During that time, bats were most common on the uninhabited islands of Amwes and Toinom (M. Marshall, pers. comm.).

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DWB observed small numbers of bats on all the islands in July 2004. The encounter rate on Namoluk Island averaged four per hour during six 45-minute walks in the least disturbed parts of the coconut-breadfruit forest on four different days. Overall, an estimate of 150–200 bats was made for the atoll. However, Maikawa Setile (pers. comm.), deputy mayor of Namoluk, claimed that bats were more abundant at certain times, especially when breadfruit was in season. He recalled seeing large numbers of bats in the settlement earlier in 2004, with as many as 50 in one tree.

+ + + Southern Mortlocks +

There are few historical accounts of flying foxes in the southern Mortlocks. J. Nason (pers. comm.) reported “scores of fruit bats... possibly 100+” on Ettal Island, Ettal Atoll, during the late 1960s.

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Wiles (1992b) reported bats on Ettal and Satawan Atolls being harvested for export in the late 1980s. A resident of Lukunor Atoll told us that he shot 50 bats on the atoll and another 150 bats on other southern Mortlock islands for this purpose during a 5-month period in 1986. These were sold to a Palauan man operating a trading boat, who provided the hunter with a .22 caliber air rifle and .410 gauge shotgun. The bats were steeped in boiling water (but not cooked), wrapped individually in plastic bags, and packed in ice prior to shipment.

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During field surveys in 2004, DWB found bats to be uncommon on Ettal Atoll and estimated 75–100 to be present. He saw only five or six bats flying in and out of several breadfruit trees in the village on Ettal Island at sunset on 30 and 31 July, and observed only one bat during several walks elsewhere on the island and none on three of the atoll’s smaller outlying islands. One resident stated that he occasionally saw as many as 20–30 bats together in breadfruit trees in the settlement.

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DWB judged bats to be common on Lekinioch Island, Lukunor Atoll, in 2004. He counted 53 bats in flight over the central taro patch and adjacent forest during 30 minutes near sunset on 2 August. No other station counts were made and no bats were seen during daytime walks among breadfruit and coconut trees in the settlement. However, the less disturbed areas of the island where bats were more likely roosting were not visited. DWB was unable to visit other islands in the atoll in 2004, but many of the people interviewed on Lekinioch, as well as former Lukunor residents living on Pohnpei at the time, reported bats on Oneop and many of the smaller islands. An overall estimate of 300–400 bats was made for the atoll.

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In 2012, DWB visited all of the islands in Lukunor Atoll except Lekinioch. Small numbers of bats were recorded on eight of the 17 islands visited, as follows: Oneop, 5 bats; Piafa, 1; Kurum, 1; Pienemon, 4; Fanamau, 1; Sapull, 6; Sopotiw, 2; and Fanafeo, 1. Seven bats were noted flying between Sapull and Sopotiw Islands. No colonies were found and no population estimate was made for the atoll.

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Surveys on Satawan Atoll in 2004 indicated that bats were common and regularly encountered, with an estimated 400–500 present. The largest numbers of bats observed during the study were on this atoll, with the greatest concentrations seen on Satawan and Ta, the two largest and southernmost islands in the Mortlocks. The encounter rate on Ta was eight per hour during walking surveys totaling 2.2 hours through breadfruit and coconut forest. The maximum number of bats observed during station counts was 81 in 25 min at Ta airport at sunset on 4 April. The largest aggregation observed during this study was 27 on Satawan Island (see Roosting below). One to 10 bats (usually less than five) were encountered per visit on other islands in the atoll, including Weito in the south; Kuttu, Orin, Pike, Mariong, Apisson, Lemasul, and Alengarik in the northwest; and Faupuker, Pononlap, and Fatikat in the east. Numbers of bats noted on Ta in 2012 matched or exceeded those recorded in 2004, and included many flying to or from Satawan Island.

+ + + Natural history of <italic><tp:taxon-name>Pteropus phaeocephalus pelagicus</tp:taxon-name></italic>. Roosting +

Pteropus phaeocephalus pelagicus characteristically roosted singly or in small, loose aggregations of 5–10 individuals, usually in the crowns of coconut and breadfruit trees in closed or nearly closed canopy forest outside of settlements. A maximum of 27 was seen together in the crowns of two adjacent coconut trees at the northern end of Satawan Island on 2 July 2004, with seven roosting along the rachis of a single palm frond and separated from each other by one or two body lengths. Bats were frequently observed hanging from tree branches and palm fronds, and occasionally clinging to the trunks of coconut trees a short distance below the crowns. Roosting bats were noted to occasionally stretch their wings and reposition themselves, or awake and relocate to another site. Some were seen crawling on the petiole bases of palm fronds and disappearing from view among the leaf axils.

+ + + Diet +

Ripe breadfruit is apparently one of the most preferred food items of Pteropus phaeocephalus pelagicus. DWB saw bats active in breadfruit trees at night on all the atolls having bats, and resident islanders invariably mentioned the fruit of breadfruit at the top of their lists of foods eaten by bats. Other foods reported by islanders were banana (Musa sp.; fruit), coconut (parts eaten unknown), papaya (Carica papaya L.; fruit), Calophyllum inophyllum L. (fruit), Crataeva speciosa Volkens (fruit), Pandanus cf. tectorius Parkinson (fruit), and Ficus tinctoria G. Forster (fruit). The only observation of bats feeding was on Weito Island, Satawan Atoll, at sunset on 3 July 2004, when three or four bats repeatedly flew in and out of the dense foliage of a pandanus tree. One flew away with a segment of ripe fruit and another dropped a segment while flying over the beach.

+ + + Flight activity +

Flying foxes were observed flying at all hours, including midday, although some daytime activity was doubtless caused by the observer’s passage. Flight activity was greatest near sunset and sunrise (Figure 7) when bats moved between day roosts and foraging sites. Fewer bats were encountered at sunrise than at sunset, suggesting that some animals returned to roosts before daylight. The direction of evening flights probably was associated with the location of food resources, often ripe breadfruit according to local residents. During April and June 2004, with the exception of a few individuals settling in nearby trees, all bats observed during evening flights at the Ta airport station moved toward the western end of the island; in July and August, 19–40% of bats flew from west to east. Interisland flights within atolls occurred regularly near sunset, but none were recorded at other times (Table 5). No bats were seen flying between atolls.

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Mean number of bats observed per 10-minute count period during (a) five early morning counts from 23 June–6 July 2004 and (b) five evening counts from 22 June–3 August 2004 at the airport on Ta Island, Satawan Atoll, Mortlock Islands.

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Numbers of bats observed flying between islands during six sunset and three sunrise counts at six different stations on Satawan Atoll, Mortlock Islands, 24 June–3 July 2004.

+ + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
StationaDateTimeNumber of batsDirection of flight
1 Fatikat 26 June1745–18150
1 Fatikat27 June0515–05450
2 Satawan (east end)2 July0500–05300
2 Satawan (east end)2 July1730–18301Fatikat to Satawan
2 Satawan (east end)3 July0430–05450
3 Satawan (west end)1 July1725–18507Ta to Satawan
4 Ta (east end)25 June1730–18300
5 Ta (west end)24 June1715–18300
6 Weito (east end)3 July1700–18358Weito to Ta
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a See Figure 1 for locations.

+ + + + + + Reproduction +

At least 50% of the bats observed on Namoluk Atoll in July 2004 appeared to be females with young. In some cases, young were evident only as a bulge beneath a female’s wing. Large volant young were occasionally seen in close proximity to or in body contact with their presumed mothers. Among five females collected on Namoluk Atoll on 21–22 July 2004, two contained single fetuses (crown-rump length 68 mm, mass 27 g, but with part of the brain case shot away; crown-rump length 42 mm) and a third had an immature male (forearm 78 mm, head-body length 110 mm) clinging to its ventral surface. Two females collected on Satawan Atoll in April 2003 each had a large young (head-body length 100, 100 mm; mass 62, 65 g) clinging to their ventral surface. Two adults were observed copulating in the crown of a coconut tree on Ta Island on 24 December 2002.

+ + + + Discussion + + Taxonomy +

Our examinations and analyses of craniometric and pelage variation in the flying foxes of Chuuk State demonstrate minor but largely consistent morphological distinctions between flying foxes of the Mortlock Islands and those from the islands of Chuuk Lagoon and Namonuito Atoll. We regard these differences as indicating no more than subspecific distinction between these two regional groupings, which we refer to Pteropus pelagicus (Pteropus phaeocephalus pelagicus and Pteropus phaeocephalus insularis, respectively). A specimen from Namonuito Atoll, the enigmatic type series of Pteropus laniger, and a series of seized bats said to originate from Chuuk State, are all best referred to Pteropus phaeocephalus insularis. The closest relative of Pteropus pelagicus is the recently extinct Pteropus tokudae of Guam, which based on its consistently smaller size, shorter and narrower rostrum, darker coloration, and considerable geographic isolation from Pteropus pelagicus, is best regarded as a distinct species (cf. Tate 1934). An overlooked nineteenth century specimen in the Berlin Museum provides an indication that the islands of Chuuk Lagoon may have originally supported a second, larger species of flying fox (i.e., in addition to Pteropus phaeocephalus insularis), similar to Pteropus pelewensis, which may now be extinct in the archipelago. The possible historical extinction of this species, and those of Pteropus tokudae on Guam and Pteropus pilosus in Palau, highlights the vulnerability of remote Pacific Pteropus populations to insular extirpation and extinction, and the importance of further clarifying studies of the taxonomic status, ecology, and current conservation standing of flying fox populations from remote Pacific archipelagos (Wiles et al. 1997, Helgen et al. 2009).

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The subspecific status of flying foxes in the northern Mortlocks remains unresolved. The nearest neighboring populations are comprised of Pteropus phaeocephalus insularis on the islands of Chuuk Lagoon 66 km to the northwest and Pteropus phaeocephalus pelagicus on Namoluk Atoll 110 km to the south. Both distances are perhaps within the flight capabilities of the species. However, based on its nearer distance and much larger land area that would support a larger bat population producing potentially more dispersing individuals, Chuuk Lagoon would appear to be a more likely source of animals colonizing Losap Atoll and Nama Island. Interisland movements of up to 119 km and corresponding genetic exchange have been reported in Pteropus mariannus in the Mariana Islands (Wiles and Glass 1990, Brown et al. 2011), but this species is substantially larger (forearm 134–154 mm), and perhaps a more powerful flier, than Pteropus pelagicus.

+ + + + Status and natural history +

The entire population of Pteropus phaeocephalus pelagicus is probably currently restricted to the four atolls comprising the southern and central Mortlock Islands, which total 10.1 km2 in size. A few additional bats may be present on Losap Atoll in the northern Mortlocks, but it is unclear which subspecies these might represent. Our study provides the first population estimate for Pteropus phaeocephalus pelagicus, with approximately925–1,200 bats present in 2004. About 75% of the population occurs on the two atolls with the largest land areas: Satawan and Lukunor. Resident islanders in the southern and central Mortlocks reported that bats were more common in the past, but that numbers appeared to have remained relatively stable in recent years. Residents also stated that bat abundance seems to fluctuate with the seasonal availability of food, especially breadfruit, but this impression likely comes from the bats’ strong attraction to villages during the peak fruiting season of breadfruit.

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Some double-counting of flying foxes may have occurred during our survey due to movements of bats between atolls. However, this problem was probably minor because our primary survey period when all six island groups in the Mortlocks were visited was limited to a 6-week period from 22 June to 5 August 2004, thus reducing the likelihood of significant inter-atoll flights.

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Recent data are lacking on the status of Pteropus phaeocephalus insularis. The main population in Chuuk Lagoon probably numbered in the mid-thousands in the early 1980s, but was reduced in abundance by commercialized hunting in the latter part of that decade (Wiles 1992b, 1992c). The only records from Namonuito Atoll are two specimens (BMNH 15.1.18.1, 15.1.18.2) collected in 1914. DWB did not sight any bats while visiting Onoun (Ulul) Island in this atoll on 3–7 January 2003, and none of the local residents queried reported seeing bats. However, no information was obtained for the islands on the eastern and northern sides of the atoll.

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Our survey efforts, including conversations with islanders, failed to confirm the presence of two other bat species in the Mortlock Islands, both of which are based on questionable early records. Thomas (1882) and Andersen (1912) listed Pteropus molossinus Temminck (originally reported under the name Pteropus breviceps; see Thomas 1887) as present in the Mortlocks, based on a single specimen collected by Kubary that supposedly originated from the island group. Pteropus molossinus is otherwise recorded only from Pohnpei and nearby Ant and Pakin Atolls (Wiles 1992d, Buden 1996a, 1996b). Our findings support Rainey and Pierson’s (1992) belief that this record is erroneous, though it is also possible that Pteropus molossinus was more widespread in the recent past and is now extinct or extremely rare in the Mortlocks. We also did not detect the insectivorous emballonurid Emballonura semicaudata sulcata Miller, which is firmly recorded only from Chuuk Lagoon’s main islands and Pohnpei (Koopman 1997). Schmeltz and Krause (1881: 298), in listing the mammals that Kubary recorded in the Mortlocks, remarked about “…noch einen von Ruk her eingefuhrten Hund, von wo auch ab und an eine Emballonura hierher verweht wird […a dog imported from Chuuk and sometimes an Emballonura blown in from that same place].” Further along in the same text, and in comparing the fauna of Nukuoro Atoll, Pohnpei State, with that of the Mortlocks, Schmeltz and Krause (1881: 331) stated, with respect to Nukuoro, “…von Säugetieren kommt nur die Ratte vor; Pteropus und Emballonura fehlen […of the mammals, there is only the rat; Pteropus and Emballonura missing.].” These unconfirmed allusions to Emballonura semicaudata in the Mortlocks are probably best treated as hypothetical or indicative of occasional vagrant occurrence.

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Atoll-dwelling populations of Pteropus are of ecological interest because of their occurrence on islands with impoverished and often highly human-altered floras (see Mueller-Dombois and Fosberg 1998). Our study of Pteropus phaeocephalus pelagicus supports the findings of others (Dolbeer et al. 1988, Wiles et al. 1991, Holmes et al. 1994) that although flying fox diets appear to comprise relatively few plant species on atolls, these islands nevertheless can support sizeable population densities approaching or exceeding 100 bats per km2. The fact that atoll habitats have experienced widespread conversion to coconut-dominated forest and that a number of cultivated fruit-producing tree species has been introduced may enhance the productivity of these islands for flying foxes (Wiles and Brooke 2009).

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Most species of Pteropus are seasonal breeders with births synchronized over a period of several months (O’Brien 1993). Contrary to this pattern, our limited observations suggest that births in Pteropus phaeocephalus pelagicus occur over a longer time frame and that females with dependent young are present for at least 8 months of the year. Two other Micronesian taxa, Pteropus yapensis in Yap and Pteropus mariannus on Guam, are among the few species of flying fox known to breed continuously (Wiles 1987b, Falanruw 1988).

+ + + Threats to populations of <italic><tp:taxon-name>Pteropus phaeocephalus pelagicus</tp:taxon-name></italic> +

With a small population size and a geographic range comprised of many small low-lying islands totaling <12 km2, flying fox populations in the Mortlock Islands are highly vulnerable to environmental changes and certain human activities. The apparent depletion or possible extirpation of bats on Losap Atoll in the northern Mortlocks during the past 60 years underscores this vulnerability.

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Sea level rise associated with climate change may represent the greatest long-term threat to Pteropus phaeocephalus pelagicus and other populations of atoll-dwelling Pteropus (Rainey 1998). One of the most recent climate models forecasts a rise in sea level of 0.57–1.10 m by 2100 and 1.84–5.39 m by 2500, although much uncertainty exists over the latter estimate (Jevrejeva et al. 2012). Rises of this magnitude certainly have the potential to submerge atolls throughout the world or greatly reduce atoll land areas and change vegetation patterns, and therefore put Pteropus phaeocephalus pelagicus at considerable risk of extinction within this or the next several centuries. Lukunor Atoll in the Mortlocks and some other Micronesian atolls experienced several extreme high tides between 2007 and 2009 that flooded islets for several hours per event (Fletcher and Richmond 2010, Keim 2010). At Lukunor Atoll, the resulting salinated soil from the initial flooding in 2007 damaged or killed 78% and 55% of the breadfruit trees on Oneop and Lekinioch islands, respectively (Keim 2010), substantially reducing an important food for Pteropus phaeocephalus pelagicus on the atoll. Despite this, some experts believe that some atoll islands may be more resilient to sea level rise than previously thought and may be able to maintain their land area through continued sediment accretion as long as geomorphic processes are not substantially altered in the future by ocean warming and acidification, changes in storm occurrence, and reductions in atoll vegetation (Woodroffe 2008, Webb and Kench 2010). However, even if atoll islands persist in the future, it is unknown what type of vegetation may remain and whether it could support populations of flying foxes.

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The negative impacts of severe cyclonic storms on Pteropus populations are well documented and occasionally include significant population declines (e.g., Cheke and Dahl 1981, Pierson et al. 1996, McConkey et al. 2004, Esselstyn et al. 2006, Wiles and Brooke 2009). The vast majority of bat mortalities occur in the aftermath of storms following depletion of food resources, loss of protected roosting sites, and in some cases increased post-storm hunting pressure. The Mortlock Islands are positioned along the southern edge of the typhoon belt present in the western North Pacific Ocean. Records since the early twentieth century suggest that individual islands in the Mortlocks experience devastating typhoons and massive damage to plant communities two to four times per century (Nason 1970, Marshall 1975, 1976, Joint Typhoon Warning Center 1980–2011, Reafsnyder 1984). However, even the largest storms rarely inflict serious habitat damage across the entire archipelago (e.g., Marshall 1979) and typically leave flying foxes and other wildlife with habitat refugia on some islands.

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In 1976, Typhoon Pamela killed most of the tree crops (e.g., breadfruit, coconut, and banana trees) on the islands of Ettal, Namoluk, Kuttu, Oneop, Moch, and Nama (Marshall 1976, 1979), including 95% of the breadfruit on Kuttu (Reafsnyder 1984) and all breadfruit, coconuts, and bananas on Ettal, where breadfruit trunks and limbs were stripped bare (M. Marshall, fide J.Nason, pers. comm.). In contrast, the islands of Losap, Ta, Satawan, Pis, and Lukunor experienced much lower losses of these trees. Ettal, Namoluk, and Kuttu were also entirely submerged by high sea surge for 15–18 hrs during this typhoon (Marshall 1979). During Typhoon Phyllis in 1958, nearly 75% of all trees at Namoluk Atoll were fully uprooted with the remaining trees being mere stumps 5–7 m tall (Davis 1959, in Marshall 1975).

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Current model projections suggest that typhoon intensity will increase, but frequency may decrease, in the western Pacific as climate change progresses over the next century (Knutson et al. 2010). Typhoon impacts on atolls may be exacerbated when combined with projected rises in sea level.

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The extent that Mortlock bat populations have been affected by severe typhoons in the past is undocumented. However, Marshall (1976) remarked that although few birds were encountered on the hardest hit islands 6–10 days after Typhoon Pamela, flying foxes appeared to survive the storm fairly well, with good numbers seen flying about searching in vain for food and shaded roosting sites. Pteropus phaeocephalus pelagicus has doubtless had a long history of exposure to intense typhoons and has been able to adjust to these periodic environmental disruptions and presumed population reductions, possibly via recruitment from surrounding islands or simply by population renewal by survivors.

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Centuries of human occupation have greatly altered the vegetation of the Mortlocks, but anthropogenic disturbance is now low despite the atoll’s high human population density. The coconut-breadfruit-pandanus forests where bats roost and feed are economically important to islanders, and they manage this resource sustainably. Cutting and clearing of the undergrowth occurs sporadically and is usually done on small, family-owned plots, but widespread cutting of forest does not occur. Additionally, a high emigration rate of Mortlockese to the larger islands of Micronesia and to overseas locations for better job opportunities (Marshall 2004) has reduced human population growth and stress on the environment.

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Overhunting has contributed to declines in flying fox populations in many areas of the Indo-Pacific region (e.g., Wiles et al. 1989, 1997, Craig et al. 1994, Brooke and Tschapka 2002, Lee et al. 2005, Wiles and Brooke 2009, Harrison et al. 2011) and may have done so in the Mortlocks in the past. Between 1986 and 1989, large numbers of Pteropus pelagicus were harvested in Chuuk State and exported to meet the demands of the commercial markets for bats on Guam (5,108 bats; Wiles 1992b) and in the Commonwealth of the Northern Mariana Islands (229 bats; Stinson et al. 1992). An unknown, but possibly sizable, number of these bats originated from the Mortlocks (Rainey 1990, Wiles 1992a, 1992b). Some of the bat exporters and/or hunters involved in this trade were from Losap, Ettal, Satawan, and Lukunor Atolls (Wiles 1992a, this study). The impacts of this harvest on bat populations in the Mortlocks were never quantified. The listing of all Micronesian species of Pteropus on Appendix I of the Convention on Trade in Endangered Species of Wild Flora and Fauna (CITES) in 1989 halted, or at least greatly reduced, commercial hunting of bats in the Federated States of Micronesia for export to the Marianas (Wiles 1992b, Wiles and Brooke 2009). It is uncertain to what extent illegal shipments of bats from Chuuk State may have occurred since then, but the amount is probably insignificant.

+

Like the vast majority of islanders from Chuuk, Pohnpei, and Kosrae states, Mortlockese almost universally disdain flying foxes as food. From the late 1960s to 1980, anthropological researchers noted that bats were not hunted or eaten on Ettal Island (Nason 1970), Namoluk Atoll (Marshall 1972), Losap Atoll (Severance 1976), Lukunor Atoll (Borthwick 1977), and Kuttu Island, Satawan Atoll (C. Reafsnyder, pers. comm.). Marshall (1972) remarked that mention of consuming bats, as done by residents of the Marianas, brought expressions of revulsion to Namoluk islanders. He also noted that bats were disliked in part because of their habit of urinating on themselves while roosting and their depredation of breadfruit, papayas, and bananas. J. Nason (pers. comm.) stated that in the Mortlockese ethnotaxonomy, bats were essentially considered as rats with wings and were never eaten. Paraphrasing a folktale told to DWB by M. Setile of Namoluk Atoll,

+

Once, long ago, the rat was envious of the bat’s wings. The rat asked the bat if he could borrow his wings for a short time just to have a brief experience of flying; he promised to return them soon. The bat allowed the rat to use his wings. But the rat lied and flew away with no intention of returning the wings. Now, what was rat is bat and the original bat is the rat.

+

Flying foxes may have been a part of the Mortlockese diet in the past, when reliance on local foods was greater and food supplies from Chuuk Lagoon and Pohnpei were not so readily available, and bats may still be utilized on occasion especially when other foods become scarce, such as after typhoons. Girschner (1912) included bats among the foods eaten by Namoluk islanders, and elders from Pis and Losap Islands told DWB of bats being eaten during the 1930s or 1940s. During our visits to the Mortlocks, no evidence was observed of bats being hunted, eaten, or killed to limit crop damage. Conversations with resident islanders throughout the central and southern Mortlocks indicated that local consumption of bats was apparently confined to a few individuals, who ate them from time to time, and to occasional visitors (largely friends and relatives) who acquired a taste for bats elsewhere.

+

Potential predators of flying foxes in the Mortlocks include four non-native species: cats (Felis catus Linnaeus), rats (Rattus exulans (Peale) and Rattus rattus complex; Wallace et al. 1972), and Pacific monitor lizards (Varanus indicus (Daudin)). Cats and rats are widely distributed among all major island groups, though not necessarily on every island. Although both may prey opportunistically on bats, Rainey (1998) noted that bats roosting in tree canopies have managed to coexist with these predators in most settings. At Namoluk Atoll, M. Setile (pers. comm.) once observed cats on the roof of a cooking shed keeping close watch on bats feeding on fruits of an adjacent Crataeva speciosa tree, but the bats kept a safe distance away. On another occasion, he saw a cat carrying the wing of a bat, but did not know how it was acquired. Monitor lizards are localized in distribution, occurring on only a few islands in the southern Mortlocks, including Lekinioch and Satawan islands (Buden 2007), where bat populations are large relative to elsewhere in the Mortlocks. Monitor lizards are opportunistic predators that consume a variety of invertebrate and smaller vertebrate prey, and are effective climbers. Flying foxes have not been reported in their diet (Dryden 1965, Losos and Greene 1988, McCoid and Witteman 1993, Dryden and Ziegler 2004, Philipp et al. 2007), but monitor lizards have been observed climbing in trees and approaching roosting Pteropus mariannus on Guam (D. Janeke, pers. comm.).

+ + + Specimens examined +

Pteropus pelagicus pelagicus

+

Mortlock Islands: Namoluk Island, Namoluk Atoll (4 skins and skulls and 2 fluid preserved adults, field numbers 12–17, plus two unnumbered fetuses and one unnumbered young, all COM collections); Satawan Island, Satawan Atoll (11 skins and 9 skulls, COM field numbers 1–11); “Mortlock Islands” (in fluid with skull extracted, BMNH 82.10.27.4 [holotype of phaeocephalus]).

+

Pteropus pelagicus insularis

+

Chuuk Lagoon islands and Namonuito Atoll: “Ruck” (skull, MNHN 1996-2112 (apparently originally A6770 fide Andersen 1912) [lectotype of insularis]; 2 skulls, ZMB 5698 and an unnumbered specimen collected in 1907; skull extracted from fluid specimen, BMNH 98.1.2.1); Uala (= Weno) (4 skins and skulls, USNM 151563–151566); Moen (= Weno) (4 skins and skulls, BYUH 248, 249, 253, 254); Namonuito (skull, BMNH 15.1.18.1); “Truk” (1 skin and skull plus 13 skulls [confiscated in Guam, probably all from Chuuk Lagoon, but possibly also outlying islands; all tentatively identified as Pteropus phaeocephalus insularis], AMNH 249954–249969); “Samoa” (emended to Caroline Islands [probably Chuuk Lagoon islands]) (skin and skull, USNM 37815 [lectotype of laniger], and skin, MCZ 7023); no locality (ANSP 6196, skull).

+

Pteropus tokudae

+

Guam (2 skins and skulls, AMNH 87117 [holotype of tokudae] and 87118, and 1 skull, AMNH 256558).

+

Pteropus cf. pelewensis

+

“Ruck” (skull, ZMB 5697); Pohnpei (skull, ZMB A4065).

+ + + + + + Acknowledgements +

We thank Phil Bruner (BYUH), Judy Chupasko and Terry McFadden (MCZ), Paula Jenkins, Richard Harbord, Roberto Portela Miguez, and Louise Tomsett (BMNH), Nancy Simmons, Eileen Westwig, and Jean Spence (AMNH), Peter Giere and Robert Asher (ZMB), Jacques Cuisin and Geraldine Veron (MNHN), Ned Gilmore (ANSP), and Darrin Lunde and Don Wilson (USNM) for loan of specimens, information on specimens in their care, for making collections available, or for other assistance during museum work. Mary Sears (Ernst Mayr Library, MCZ) kindly provided photocopies of library materials from special collections, including a scanned image of Pteropus phaeocephalus from the plate that accompanied the original description. Doug Kelly (COM) assembled the composite illustration in Figure 3. Lauren Helgen assisted with specimen photography and in preparing figures (Figures 2, 4, and 6). We also thank Bernhard Hausdorf, Ronald Janssen, and Katrin Schniebs for responding to queries concerning J. Kubary in the Mortlock Islands, Vladimir Loskot for his search of collection records in the Russian Academy of Sciences, and Mark Borthwick, James Nason, Charles Reafsnyder, Craig Severance, Mac Marshall, and Harley Manner for sharing their recollections on the status of bats in the Mortlocks. Paula Jenkins commented on portions of a preliminary draft of the manuscript.

+

We thank Al Gardner, Richard Hoffman, Tony Hutson, Paula Jenkins, Tim McCarthy, Storrs Olson, and Van Wallach for sharing their taxonomic views and knowledge of nomenclatural procedures, and Micaela Jemison for helpful suggestions. Paul Clark, R. Clouse, Doug Craig, A. Gardner, Claudette Lucand, S. Olson, Leslie Overstreet, and Geraldine Veron furnished information on the Voyage au Pole Sud.

+

We are especially grateful to the many people of the Mortlock Islands who assisted DWB in many different ways during his visits, particularly his island hosts, including Mitaro Josa on Nama, Maikawa Setile on Namoluk, Benjamin Johnna on Ettal, Richard Carlos on Satawan, Sander Mark on Kuttu, and Theodor Emuch on Moch. A special note of thanks is extended to Obed Mwarluk on Ta, who provided food and lodging during DWB’s many visits to Satawan Atoll and assisted with the logistics of travel and accommodations elsewhere in the Mortlocks.

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\ No newline at end of file From 416ea4309f2b14b47b7ff5a8d15b3c58a01a4d0b Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Mon, 24 Mar 2014 08:06:37 +0000 Subject: [PATCH 16/18] Renamed 10.3897/zookeys.345.5840 --- 10.3897:zookeys.345.5840.xml => 10.3897_zookeys.345.5840.xml | 0 1 file changed, 0 insertions(+), 0 deletions(-) rename 10.3897:zookeys.345.5840.xml => 10.3897_zookeys.345.5840.xml (100%) diff --git a/10.3897:zookeys.345.5840.xml b/10.3897_zookeys.345.5840.xml similarity index 100% rename from 10.3897:zookeys.345.5840.xml rename to 10.3897_zookeys.345.5840.xml From b76a2338bbd6ff0f9ffb44b5439dc600cee03ab3 Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Thu, 1 May 2014 08:35:29 +0100 Subject: [PATCH 17/18] Edited some references --- 10.3897_zookeys.183.3073.xml | 4 ++-- 1 file changed, 2 insertions(+), 2 deletions(-) diff --git a/10.3897_zookeys.183.3073.xml b/10.3897_zookeys.183.3073.xml index 150862d..13a6c39 100644 --- a/10.3897_zookeys.183.3073.xml +++ b/10.3897_zookeys.183.3073.xml @@ -220,10 +220,10 @@ Banner AH (1953) The Crangonidae, or snapping shrimp, of Hawaii. Pacific Science 7: 3-147. Banner DMBanner AH (1982). The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum 34: 1-357. doi: 10.3853/j.0067-1975.34.1982.434 Boone L (1930) Scientific results of the cruises of the yachts “Eagle” and “Ara”, 1921–1928, William K. Vanderbilt, commanding. Crustacea: Anomura, Macrura, Schizopoda, Isopoda, Amphipoda, Mysidacea, Cirripedia, and Copepoda. Bulletin of the Vanderbilt Marine Museum 3: 1-221. - Burukovsky RN (1990) Shrimps from the Sala-y-Gomez and Nazca Ridges [in Russian]. Trudy Instituta Okeanologii Akademii Nauk SSSR 124: 187–217. + Burukovsky RN (1990) Shrimps from the Sala-y-Gomez and Nazca Ridges [in Russian]. Trudy Instituta Okeanologii Akademii Nauk SSSR 124: 187–217. Chace FAJr (1966) Decapod crustaceans from St. Helena, South Atlantic.  Proceedings of the United States National Museum 118: 622-662. Chace FAJr (1972) The shrimps of the Smithsonian–Bredin Caribbean expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology 98: 1-179. - Christoffersen ML (1979) Campagne de la Calypso au large des côtes Atlantiques de l’Amerique du Sud (1961–1962). I. Decapod Crustacea: Alpheoida. Annales de l’Institut Océanographique 55(Suppl.): 297–377. + Christoffersen ML (1979) Campagne de la Calypso au large des côtes Atlantiques de l’Amerique du Sud (1961–1962). I. Decapod Crustacea: Alpheoida. Annales de l’Institut Océanographique 55(Suppl.): 297–377. Christoffersen ML (1998) Malacostraca. Eucarida. Caridea, Crangonoidea and Alpheoidea (except Glyphocrangonidae and Crangonidae). In: Young PS (Ed) Catalogue of Crustacea of Brazil. Museu Nacional, Rio de Janeiro: 351-372. Coutière H (1897) Note sur quelques Alphées nouveaux. Bulletin du Muséum d’Histoire Naturelle, serie 3 7: 303–306. Coutière H (1905) Les Alpheidae. In: Gardiner JS The Fauna and Geography of the Maldive and Laccadive Archipelagoes. Being the account of the work carried on and of the Collections made by an Expedition during the years 1899 and 1900. University Press, Cambridge, 852–921, pls. 70–87. From 4b111b1ad51709c02cac6926ec100816579dbacc Mon Sep 17 00:00:00 2001 From: Roderic Page Date: Thu, 1 May 2014 08:43:20 +0100 Subject: [PATCH 18/18] Deleted extra tag --- 10.3897_zookeys.183.3073.xml | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/10.3897_zookeys.183.3073.xml b/10.3897_zookeys.183.3073.xml index 13a6c39..c1388f2 100644 --- a/10.3897_zookeys.183.3073.xml +++ b/10.3897_zookeys.183.3073.xml @@ -223,7 +223,7 @@ Burukovsky RN (1990) Shrimps from the Sala-y-Gomez and Nazca Ridges [in Russian]. Trudy Instituta Okeanologii Akademii Nauk SSSR 124: 187–217. Chace FAJr (1966) Decapod crustaceans from St. Helena, South Atlantic.  Proceedings of the United States National Museum 118: 622-662. Chace FAJr (1972) The shrimps of the Smithsonian–Bredin Caribbean expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology 98: 1-179. - Christoffersen ML (1979) Campagne de la Calypso au large des côtes Atlantiques de l’Amerique du Sud (1961–1962). I. Decapod Crustacea: Alpheoida. Annales de l’Institut Océanographique 55(Suppl.): 297–377. + Christoffersen ML (1979) Campagne de la Calypso au large des côtes Atlantiques de l’Amerique du Sud (1961–1962). I. Decapod Crustacea: Alpheoida. Annales de l’Institut Océanographique 55(Suppl.): 297–377. Christoffersen ML (1998) Malacostraca. Eucarida. Caridea, Crangonoidea and Alpheoidea (except Glyphocrangonidae and Crangonidae). In: Young PS (Ed) Catalogue of Crustacea of Brazil. Museu Nacional, Rio de Janeiro: 351-372. Coutière H (1897) Note sur quelques Alphées nouveaux. Bulletin du Muséum d’Histoire Naturelle, serie 3 7: 303–306. Coutière H (1905) Les Alpheidae. In: Gardiner JS The Fauna and Geography of the Maldive and Laccadive Archipelagoes. Being the account of the work carried on and of the Collections made by an Expedition during the years 1899 and 1900. University Press, Cambridge, 852–921, pls. 70–87.