A catalogue of the collections of Mexican amber at the Natural History Museum, London and National Museums Scotland, Edinburgh, UK

Ross, Andrew J.; Mellish, Claire J.T.; Crighton, Bill; York, Peter V.; Ross, Andrew J.; Mellish, Claire J.T.; Crighton, Bill; York, Peter V.

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Boletín de la Sociedad Geológica Mexicana

Print version ISSN 1405-3322

Bol. Soc. Geol. Mex vol.68 n.1 Ciudad de México Apr. 2016

Articles

A catalogue of the collections of Mexican amber at the Natural History Museum, London and National Museums Scotland, Edinburgh, UK

Catálogo de las colecciones de ámbar mexicano en el Museo de Historia Natural de Londres y en los Museos Nacionales de Escocia en Edimburgo, Reino Unido

Andrew J. Ross¹²^*

Claire J.T. Mellish²

Bill Crighton¹

Peter V. York³

^¹ Department of Natural Sciences, National Museum of Scotland, Chambers St., Edinburgh, EH1 1JF.

^² Department of Earth Sciences, Natural History Museum, Cromwell Road, London, SW7 5BD.

^³ Department of Life Sciences, Natural History Museum, Cromwell Road, London, SW7 5BD.

Abstract:

A catalogue is here provided of the pieces of Mexican amber with inclusions in the collections of the Natural History Museum, London, and National Museums Scotland, Edinburgh, both in the United Kingdom. There are 32 pieces in the Natural History Museum and 101 pieces in National Museums Scotland which contain a combined total of 557 arthropod and 13 plant inclusions. Four orders and 11 families of arthropods are additional to a taxonomic list of Mexican amber arthropods published in 2010.

Keywords: Mexican amber; inclusions; arthropods; London; Edinburgh

Resumen:

Presentamos aquí el catálogo de piezas de ámbar de México con inclusiones encontrados en las colecciones del Museo de Historia Natural de Londres y del Museo Nacional de Escocia en Edimburgo, ambos en el Reino Unido. En el Natural History Museum encontramos 32 piezas, mientras que, en el Museo Nacional de Escocia, encontramos un total de 101 piezas. Se incluye un total de 557 artrópodos y 13 inclusiones de plantas. Cuatro órdenes y 11 familias de artrópodos son adicionales a la lista taxonómica mexicana de artrópodos en ámbar publicada en el 2010.

Palabras clave: ámbar de México; inclusiones; artrópodos; Londres; Edimburgo

1. Introduction

For the past 20 years the Natural History Museum in London (NHM) and National Museums Scotland in Edinburgh (NMS) have been acquiring specimens of Mexican amber with inclusions by purchase or donation. This paper comprises a list of the specimens to make the wider scientific community aware of this material and to hopefully generate research on the specimens.

There are currently 32 pieces of Mexican amber with inclusions in the collection of the NHM and 101 pieces in NMS. Many of these pieces contain several inclusions and contain a combined total of 557 arthropods and 13 plant remains (although one may be fungal).

Most of the specimens have not been studied in detail so the identifications were predominantly undertaken by AJR. Over the years various visiting scientists have improved some of the IDs and a few new taxa have been described.

2. Catalogue

The catalogue below is organised by major taxonomic group (in capitals), then by order and family where possible. Order and family names of arthropods given in bold were not included in ^{Solórzano Kraemer (2010)}, of which there are four additional orders and eleven additional families listed here. For those where a family ID has not yet been provided then 'Undetermined' is used though some of these specimens may be fragmentary or poorly preserved and so are indeterminate. Specimen numbers pre-fixed by a question mark show that the identification is tentative. The total number of inclusions is given in brackets after the taxonomic name, excluding trace fossils. Further identifications and type details are also provided.

Twenty four specimens are figured here, including previously described holotypes. Fourteen of these specimens have not been previously figured. The corresponding figure number is given in brackets, in bold, after the specimen number. All scale bars represent 1 mm.

The specimens at the NHM are pre-fixed by II (for insects), IA (for arachnids) or V (for plants). The main three or four-digit number is the number of the piece of amber; the number in brackets denotes the number for the taxon where more than one taxon occur in the one piece; if there is more than one specimen of that taxon in the amber then the quantity is given in brackets, e.g . II.2132(2) (x2) denotes amber piece with insects no. 2132, taxon no. 2 and x2 specimens of that taxon.

The specimens at the NMS are prefixed with G followed by the year of acquisition, the batch number, the number of the amber piece, the number of the taxon if there is more than one, and the quantity of that taxon is given in brackets if more than one specimen exists, e.g . G.2004.6.10.1 (x3) denotes year of acquisition 2004, batch no. 6, amber piece no. 10, taxon no. 1 and x3 specimens of that taxon. The specimens are kept at the National Museums Collection Centre in Edinburgh, which is part of National Museums Scotland.

The catalogue was compiled by AJR and CJTM, with photographs of NMS specimens by BC (Copyright NMS) and NHM specimens by PVY (Copyright NHM), unless otherwise stated in the figure caption.

ARACHNIDA (52)

Acari (3):

Undetermined (3):
• G.2004.50.3.3, G.2008.34.14.7, G.2011.31.8.3

Araneae (47):

Linyphiidae (1):
• G.2004.50.13
Mimetidae (1):
• G.2004.50.11
Miturgidae (2):
• G.2004.6.2.1, G.2004.50.8.1
Mygalomorpha undetermined (1):
• G.2004.50.9.1
Pholcidae(1):
• Modisimus sp. G.2004.6.4.1 (Figure 1)

Figure 1 Daddy long-legs spider (Araneae: Pholcidae), Modisimus sp., NMS G.2004.6.4.1.

Salticidae (5):
• G.2004.50.3.1, G.2004.50.4.1, G.2004.50.5.1, G.2004.50.12.1, G.2005.147.9
Theraphosidae (1):
• Figured ^{Dunlop et al. (2008}, pp. 13-14, Figs 2-9) G.2004.6.1.1
Trochanteriidae (4?):
• Veteratorsp. G.2004.50.1.1, ?G.2004.50.2.1, ?G.2004.50.10.1 (x2)
Undetermined (31):
• IA.61 (with II.2159 and V.68584), IA.62 (with II.2163 and V.68586), G.1998.54.3.1, G.2004.6.3.1 (x2), G.2004.6.10.1 (x3), G.2004.7.1.1, G.2004.50.3.2, G.2004.50.6, G.2004.50.7.1, G.2005.147.6.1, G.2005.147.7.1, G.2005.147.8, G.2005.147.10, G.2005.147.11.1, G.2006.42.4, G.2006.42.5.1, G.2006.42.7.5, G.2007.67.1.1, G.2007.67.2, G.2007.67.3, G.2007.67.4, G.2007.67.5.4, G.2007.67.14.6, G.2008.34.3.4, G.2008.34.9.3, G.2008.34.13, G.2008.34.14.2, G.2014.50.6.2

Pseudoscorpiones (2):

Atemnidae(1):
• G.2004.6.5.1
Chernetidae (1):
• Figured ^{Ross and Sheridan (2013}, p. 54, Figure 2) G.2011.31.2.1

MYRIAPODA (5):

Chilopoda (3):

Geophilomorpha(2):
• G.2007.67.10.1, Figured ^{Ross and Sheridan (2013}, p. 55, Fig. 9) G.2005.147.1.1 (Figure 2)

Figure 2 Centipede (Chilopoda: Geophilomorpha), NMS G.2005.147.1.1.

Scutigeromorpha(1):
• G.2014.50.2 (Figure 3)

Figure 3 House centipede (Chilopoda: Scutigeromorpha), NMS G.2014.50.2.

Diplopoda (2):

Undetermined (2):
• G.2005.147.2, G.2006.42.2

CRUSTACEA (1):

Isopoda (1):

Undetermined (1):
• G.2008.34.14.1

HEXAPODA (496):

Collembola (11):

Undetermined (11):
• II.1863(4), II.2164(2), II.3084(7), G.2004.6.10.3, G.2006.42.3.2, G.2006.42.5.2, G.2008.34.11.5 (x2), G.2014.50.6.3 (x2), G.2014.50.7.3

Archaeognatha (2):

Machilidae (2):
• G.2004.6.6.1, G.2005.147.3.2

Ephemeroptera (3):

Baetidae (1):
• II.2159(1) (Figure 4)

Figure 4 Mayfly (Ephemeroptera: Baetidae), NHM II.2159(1). Photo taken by Harry Taylor (NHM).

Heptageniidae(2):
• Maccaffertium annae ^{Macadam and Ross, 2016}, Holotype G.2008.34.3.1, Paratype G.2008.34.3.2

Odonata(1):

Coenagrionidae(1):
• Neoerythrommasp. Figured Ross, Coutiño José and Nel, 2016 G.2008.34.1.1
Blattodea (13):
Ectobiidae (=Blattellidae) (8):
• II.1863(1), II.2165, II.2167(1), G.2014.50.7.1, G.2014.50.8.1• Anaplecta sp. II.2168(1), G.2011.31.10, Figured ^{Ross and Sheridan (2013}, p. 55, Fig. 8) G.2006.42.1 (Figure 5)

Figure 5 Cockroach (Blattodea: Ectobiidae), Anaplecta sp., NMS G.2006.42.1.

Undetermined (5):
• II.1862, II.2159(6), G.2004.50.3.4, G.2008.34.11.1, G.2014.50.9.1

Dermaptera(1):

Diplatyidae(1):
• Haplodiplatys crightoni^{Ross and Engel, 2013}, Holotype G.2011.31.1.1 (Figure 6)

Figure 6 Earwig (Dermaptera: Diplatyidae), Holotype of Haplodiplatys crightoni ^{Ross and Engel, 2013}, NMS G.2011.31.1.1.

Isoptera (29):

Mastotermitidae (6):
• Mastotermessp. II.2159(3), G.2005.147.7.2, G.2008.34.6 (x2), G.2014.50.3.1, Figured ^{Ross and Sheridan (2013}, p. 54, Fig. 3) G.2005.147.4.1
Undetermined (23):
• II.2132(2) (x2), II.2133(1), II.2158(2), II.2160(1), II.2166(2) (x7), G.1998.38.6.1 (x4), G.2000.63.3, G.2005.147.4.2, G.2005.147.5, G.2006.42.6, G.2007.67.11.3, G.2007.67.14.1, G.2008.34.14.3

Orthoptera (2):

Grylloidea undetermined (2):
• II.2166(1) (Figure 7), G.2014.50.1 (Figure 8)

Figure 7 Cricket (Orthoptera: Grylloidea), NHM II.2166(1).

Figure 8 Cricket (Orthoptera: Grylloidea), NMS G.2014.50.1.

Hemiptera (49):

Aetalionidae(1):
• II.2127(1)
Aleyrodidae (2):
• G.2004.6.10.5, G.2007.67.13.3
Cicadellidae (4):
• G.2004.50.3.5, G.2006.42.7.9, G.2007.67.11.2, Figured ^{Ross and Sheridan (2013}, p. 56, Fig. 13) G.2008.34.5.1
Cixiidae (6):
• II.2132(1), G.2000.63.1, G.2006.42.7.3, G.2011.31.1.5, G.2011.31.5 (Figure 9), G.2014.50.5.1

Figure 9 Planthopper (Hemiptera: Cixiidae), NMS G.2011.31.5.

Coccoidea undetermined (3?):
• G.2004.50.3.6, G.2008.34.3.8, ?G.2008.34.10.3
Derbidae (1):
• Cedusa baylissae^{Szwedo and Ross, 2003}, Holotype II.2154 (Figure 10)

Figure 10 Planthopper (Hemiptera: Derbidae), Holotype of Cedusa baylissae ^{Szwedo and Ross, 2003}, NHM II.2154.

Fulgoroidea undetermined (14):
• II.2127(2), II.3084(5), G.2004.6.1.2 (x3), G.2004.50.9.3, G.2005.147.6.2, G.2006.42.7.2, G.2008.34.3.3, G.2008.34.10.1, G.2011.31.4.2 (x2), G.2014.50.5.2, Figured ^{Ross (1998}, Fig. 70; 2010, Fig. 80) II.1869
Heteroptera undetermined (2):
• II.2133(10), G.2007.67.11.1
Homoptera undetermined (10?):
• II.2168(6), G.2004.6.1.3 (x4), G.2004.6.4.2, G.2004.50.2.2, ?G.2006.42.5.5, G.2008.34.14.6, G.2014.50.5.7
Kinnaridae(1):
• G.2014.50.6.1 (Figure 11)

Figure 11 Planthopper (Hemiptera: Kinnaridae), NMS G.2014.50.6.1.

Nogodinidae (1):
• Tonacatecutlius gibsoni^{Stroiński and Szwedo, 2008}, Holotype II.2131(1) (Figure 12)

Figure 12 Planthopper (Hemiptera: Nogodinidae), Holotype of Tonacatecutlius gibsoni ^{Stroiński and Szwedo, 2008}, NHM II.2131(1).

• Figured ^{Ross and Sheridan (2013}, p. 56, Fig. 14) G.2008.34.7 (Figure 13)

Figure 13 Assassin bug (Hemiptera: Reduviidae), NMS G.2008.34.7.

Tropiduchidae(1):
• Figured ^{Ross and Sheridan (2013}, p. 56, Fig. 16) G.2011.31.4.1
Undetermined (2):
• G.2011.31.2.4 (x2)

Psocoptera (13):

Undetermined (13):
• II.1863(3), II.1865(1), II.2132(3), II.3084(6), G.1998.38.6.2, G.2004.6.10.4, G.2004.7.1.2, G.2004.50.3.7 (x2), G.2004.50.7.2, G.2007.67.14.5, G.2014.50.4.2, G.2014.50.5.5

Thysanoptera (5):

Undetermined (5):
• II.2133(2), II.3084(8), G.2007.67.14.10, G.2011.31.7.3 (x2)

Coleoptera (44):

Anthribidae (1):
• Figured ^{Ross and Sheridan (2013}, p. 55, Fig. 6) G.2004.6.8.1 (Figure 14)

Figure 14 Fungus weevil (Coleoptera: Anthribidae), NMS G.2004.6.8.1.

Cerambycidae (2):
• G.2007.67.12.1, Figured ^{Ross and Sheridan (2013}, p. 56, Fig. 17) G.2008.34.2 (Figure 15)

Figure 15 Long-horn beetle (Coleoptera: Cerambycidae), NMS G.2008.34.2.

Curculionidae (2):
• Platypodinae G.2007.67.12.2 (x2)
Curculionoidea undetermined (1):
• II.1863(2)
Dytiscidae (9?):
• Figured ^{Ross and Sheridan (2013}, p. 54, Fig. 1) ?G.2006.42.3.1 (x9)
Elateroidea undetermined (3):
• II.2164(1), II.3083(1) (Figure 16), G.2008.34.9.1

Figure 16 Click beetle (Coleoptera: Elateroidea), NHM II.3083(1).

Staphylinidae (4):
• G.1998.38.6.3, G.2004.50.7.3, G.2007.67.12.3, G.2014.50.9.2
Undetermined (22):
• II.2133(3) (x2), G.1998.38.3.2, G.2004.6.1.4, G.2004.6.9, G.2004.6.10.7 (x6), G.2004.50.3.9, G.2004.50.4.2, G.2004.50.10.2, G.2005.147.9.3, G.2006.42.7.8, G.2007.67.7.1, G.2007.67.12.4 (x2), G.2011.31.2.3, G.2014.50.6.4, G.2014.50.8.3

Diptera (193):

Cecidomyiidae (63):
• II.1864(2) (x2), II.1865(4), II.2132(5), II.2133(8) (x4), II.2160(2) (x2), II.2168(3) (x4), II.3083(4) (x2), II.3084(3), G.2004.6.1.7, G.2004.6.2.2, G.2004.6.3.3, G.2004.6.10.9 (x4), G.2004.50.3.11 (x6), G.2005.147.9.2, G.2006.42.7.7 (x5), G.2007.67.10.2, G.2007.67.12.6, G.2007.67.14.3, G.2008.34.1.2, G.2008.34.11.3, G.2008.34.14.5, G.2014.50.5.6 (x2), G.2014.50.7.4, G.2014.50.8.5• Lestremia sp. G.2004.50.1.2 (x6), G.2005.147.6.3, G.2006.42.5.4, G.2007.67.5.3 (x8), G.2011.31.7.2
Ceratopogonidae (5):
• II.1864(3), II.2159(5), II.2168(5). G.2004.6.1.6, G.2014.50.4.6
Chironomidae (25):
• II.1863(6), II.1864(3), II.1865(3), II.1866(2), II.2133(7), II.2166(4), II.2167(3) (x2), II.2168(2) (x4), II.3083(2) (x2), G.2006.42.7.6, G.2007.67.6.3, G.2008.34.3.7 (x2), G.2008.34.11.4, G.2014.50.4.4 (x3), G.2014.50.8.4, G.2014.50.9.3 (x2)
Culicidae (2):
• G.2004.6.1.5 (x2)
Cyclorrapha undetermined (2):
• II.2166(5), G.2007.67.10.3
Dolichopodidae (9):
• II.1871, II.2168(9), G.1998.38.6.5, G.2004.6.3.2, G.2004.6.10.10, G.2007.67.14.8, G.2011.31.9.1 (x3)
Drosophilidae (1):
• II.1864(5)
Keroplatidae (2):
• G.2004.50.3.10, G.2007.67.14.2
Limoniidae (2):
• G.2000.63.2• Toxorhina sp. II.1870
Mycetophilidae (2):
• II.1866(1) (Figure 17), G.2004.7.1.3

Figure 17 Fungus gnat (Diptera: Mycetophilidae), NHM II.1866(1).

Nematocera undetermined (7):
• G.2004.7.1.4 (x2), G.2007.67.12.5, G.2007.67.14.9 (x2), G.2014.50.9.4 (x2)
Phoridae (22):
• II.894(1), II.895(1), II.2133(9), II. 2159(2), II.2163 (x11), II.3084(4), G.2004.50.3.12 (x2), G.2004.50.5.2 G.2008.34.5.2 (x3)
Psychodidae (25):
• II.894(2), II.1865(2), II.2133(12), II.2160(3), II.2168(8), G.2004.6.4.3 (x3), G.2004.6.10.8 (x6), G.2006.42.5.3 (x2), G.2007.67.14.7, G.2008.34.11.2, G.2008.34.14.4, G.2011.31.1.3, G.2014.50.4.3 (x4)
• Phlebotominae II.1863(7)
Rhagionidae(1):
• Figured ^{Ross and Sheridan (2013}, p. 55, Fig. 4) G.2008.34.4
Scatopsidae (6):
• II.1867(1), II.3083(3) (x2), G.1998.38.6.4, G.2007.67.1.2, G.2014.50.4.5
Sciaridae (7):
• II.1865(5), II.2133(11), II.2168(4), G.2004.6.10.11, G.2007.67.6.2, G.2007.67.14.4, G.2014.50.3.2
Sciaroidea undetermined (2):
• II.2166(3), G.2007.67.13.2
Stratiomyidae (1):
• G.2008.34.12 (Figure 18)

Figure 18 Soldier fly larva (Diptera: Stratiomyidae), NMS G.2008.34.12.

Tephritidae(1):
• II.2169 (Figure 19)

Figure 19 Fruit fly (Diptera: Tephritidae), NHM II.2169.

Therevidae(1):
• Peratrimera mexicana^{Hauser and Irwin, 2005}, Holotype II.2158(1) (Figure 20)

Figure 20 Stiletto fly (Diptera: Therevidae), Holotype of Peratrimera mexicana ^{Hauser and Irwin, 2005}, NHM II.2158(1).

Tipuloidea undetermined (3)
• G.2005.147.3.3, G.2007.67.6.1, G.2008.34.9.2
Undetermined (4)
• G.2004.6.10.12 (x2), G.2011.31.4.3, G.2011.31.7.1

Hymenoptera (97):

Apidae (14):
• Meliponini II.3084(1) (x3), G.2004.50.1.3 (x6), G.2004.50.2.3, G.2005.147.11.2, G.2011.31.9.2 (x2), G.2014.50.5.3
Apocrita undetermined (3):
• G.2007.67.5.2, G.2014.50.4.7 (x2)
Bethylidae (1):
• II.2162 (Figure 21)

Figure 21 Bethylid wasp (Hymenoptera: Bethylidae), NHM II.2162.

Braconidae (2):
• II.1868 (Figure 22), G.2011.31.1.2

Figure 22 Braconid wasp (Hymenoptera: Braconidae), NHM II.1868. Photo taken by Harry Taylor (NHM).

Chalcidoidea undetermined (3):
• II.2168(7), G.2004.50.3.14, G.2008.34.3.5
Formicidae (34):
• II.895(2), II.1867(2), II.2131(2), II.2159(4), II.3084(2) (x3), G.1998.38.3.3, G.1998.38.6.6, G.1998.54.3.3, G.2004.6.6.2, G.2004.6.8.2, G.2004.6.10.2 (x4), G.2004.7.1.6 (x3), G.2004.50.4.3, G.2004.50.9.4, G.2004.50.12.2, G.2005.147.1.2, G.2007.67.9 (x5), G.2008.34.1.4, G.2008.34.10.2, G.2011.31.2.2, G.2011.31.8.2, G.2014.50.5.4, G.2014.50.7.2 Ichneumonoidea undetermined (1):
• Figured ^{Ross and Sheridan (2013}, front cover, p. 6) G.1998.54.3.2
Mymaridae (6):
• II.2133(5) (x2), G.2004.6.3.4, G.2004.7.1.7, G.2004.50.3.15, G.2004.50.7.5
Platygastroidea undetermined (1):
• II.2133(4)
Scelionidae (1?):
• ?II.1872
Parasitica undetermined (31):
• II.1863(5), II.2132(4), II.2133(6), II.2158(3), II.2167(2), II.3083(5), G.2004.6.10.13 (x4), G.2004.7.1.5 (x4), G.2004.50.1.4 (x2), G.2004.50.3.13, G.2004.50.7.4, G.2005.147.3.4, G.2005.147.9.4, G.2006.42.7.4 (x7), G.2008.34.1.3, G.2008.34.3.6, G.2011.31.1.4 (x2)

Lepidoptera (10):

Undetermined (10):
• II.2161, G.2004.6.10.6, G.2004.50.3.8, G.2004.50.9.2, G.2006.42.7.1, G.2007.67.5.1, G.2007.67.13.1, G.2011.31.6, G.2014.50.8.2,• Figured ^{Ross and Sheridan (2013}, p. 56, Fig. 18) G.2008.34.8

Trichoptera (10):

Glossosomatidae (1):
• II.1864 (6)
Hydropsychidae (4):
• Plectropsyche alvarezi^{Wichard, Solórzano Kraemer and Luer, 2006} (det. Wilfried Wichard) II.1864(1) (x4) (Figure 23)

Figure 23 Caddisfly (Trichoptera: Hydropsychidae), Plectropsyche alvarezi ^{Wichard, Solórzano Kraemer and Luer, 2006}, NHM II.1864(1).

Undetermined (5):
• G.2008.34.3.9, G.2014.50.4.1 (x4)

Undetermined larvae (16):

• G.2004.6.5.2 (x14), G.2007.67.7.2, G.2007.67.8

Insect coprolites:

• G.2007.67.15

MOLLUSCA:

Bivalvia:

Pholadidae borings
• G.2011.31.14

PLANTAE (13?):

Marchantiophyta (1):

Jungermanniales (1):
Lejeuneaceae (1?):
• Figured ^{Ross and Sheridan (2013}, p. 55, Fig. 11) ?G.1998.38.3.1

Angiospermae (8):

Ericales (1):
Ericaceae (1?):
• Figured ^{Ross and Sheridan (2013}, p. 55, Fig. 7) ?G.2004.6.7
Undetermined (7):
• V.68584 (with IA.61 and II.2159), V.68585 (with II.3084), V.68586 (with IA.62 and II.2163), G.2004.50.5.3, G.2005.147.3.1, G.2011.31.8.1, Figured ^{Solórzano Kraemer (2010}, Fig. 8A), ^{Ross and Sheridan (2013}, p. 56, Fig. 12) G.2007.67.16

Coniferae (1):

Undetermined (1):
• G.2008.34.16

Undetermined (3):

• G.2008.34.17, G.2011.31.3, G.2014.50.7.5

Plantae or Fungi undetermined (1):

• Figured ^{Ross and Sheridan (2013}, p. 56, Fig. 15) G.2008.34.15 (Figure 24)

Figure 24 Undetermined plant or fungal structure shedding spores, NMS G.2008.34.15.

Acknowledgements

Many thanks to Harry Taylor (NHM) for photographing the mayfly in Figure 4, and to Vicen Carrio for writing the Spanish translation of the abstract.

References

Dunlop, J.A., Harms, D., Penney, D., 2008, A fossil tarantula (Araneae: Theraphosidae) from Miocene Chiapas amber, Mexico: Revista Ibérica de Aracnología, 15, 9-17. [ Links ]

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Received: November 14, 2014; Revised: March 27, 2015; Accepted: April 05, 2015

^* a.ross@nms.ac.uk

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