Orthoptera: Acrididae: Oedipodinae
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Zoological Studies 47(4): 495-506 (2008) A Review of the Canarian Sphingonotini with Description of a New Species from Fuerteventura (Orthoptera: Acrididae: Oedipodinae) Axel Hochkirch1,* and Martin Husemann2 1University of Trier, Department VI, Biogeography Group, Am Wissenschaftspark 25-27, Trier, D-54296, Germany 2University of Osnabrück, Department of Biology/Chemistry, Division of Ecology, Barbarastr 11, Osnabrück, D-49076, Germany (Accepted November 19, 2007) Axel Hochkirch and Martin Husemann (2008) A review of the Canarian Sphingonotini with description of a new species from Fuerteventura (Orthoptera: Acrididae: Oedipodinae). Zoological Studies 47(4): 495-506. The Canarian representatives of the tribe Sphingonotini (Orthoptera: Acrididae: Oedipodinae) are revised, and a molecular phylogeny based on mitochondrial (mt)DNA sequences (ND5) is presented. The genera Wernerella, Pseudosphingonotus, and Neosphingonotus are synonymized with Sphingonotus. A new grasshopper species from Fuerteventura, Sphingonotus fuerteventurae sp. nov., is described and compared with the Canarian Sphingonotini and some closely related West-Mediterranean species. Some information on the distribution and ecology of the new species is given, and a key to the Canarian species of Sphingonotus is presented. http://zoolstud.sinica.edu.tw/Journals/47.4/495.pdf Key words: Taxonomy, Canary Is., Cryptic species, Sphingonotini, Sphingonotus. The Canarian archipelago is one of the of some species remains a matter of controversy global hot spots of endemism, with 27% of its (Bland et al. 1996, Hochkirch 2003). This has been native flora and 50% of the terrestrial invertebrate particularly true for the 3 closely related species fauna endemic (Juan et al. 2000). The insect S. rubescens (Walker, 1870), S. caerulans fauna comprises approximately 2200 endemic (Linnaeus, 1767) and S. corsicus Chopard, species (Oromí and Baéz 2005). About 86 1923 (see also Bland et al. 1996 for discussion). orthopteran (sensu stricto) species are known to Although Uvarov (1923) long ago stressed occur on the archipelago, including 33 endemics morphological differences among these species, (Bland et al. 1996, Bland 2001, López et al. 2005, uncertainty with their identification still remains, Pfau and Pfau 2007). Although the Canary Is. is a and even experienced taxonomists have argued major travel destination of European tourists, their that these species are difficult to distinguish faunistic exploration is far from complete. Four (e.g., Willemse 1985). Recent studies show that new species and subspecies of Orthoptera have S. rubescens can easily be morphometrically been described during the last decade (Bland and distinguished from other species of the S. caerulans Gangwere 1998, Bland 2001, López et al. 2005, group (Defaut 2003) and by its characteristic song Pfau and Pfau 2007), and some species have been (Bland 1985, Husemann and Hochkirch 2008). recorded for the 1st time from single islands (e.g., Sphingonotus rubescens has a longer and more- Hochkirch 2003, Husemann and Hochkirch 2008). slender body and longer wings than S. caerulans. The genus Sphingonotus Fieber, 1852 and its Its hindwings are hyaline with blackish vannal relatives (Pseudosphingonotus Shumakov, 1963 veins, whereas S. caerulans and S. corsicus have and Wernerella Karny, 1907) are particularly well bluish hindwings and a denser reticulation of the represented on the Canary Is., although the identity tegmina. At present, no confirmed records of * To whom correspondence and reprint requests should be addressed. Tel: 49-651-2014692. Fax: 49-651-2013851. E-mail:[email protected] 495 496 Zoological Studies 47(4): 495-506 (2008) S. caerulans or S. corsicus exist from the Canary A mitochondrial gene fragment (ND5, 1059 bp) Is. was amplified and sequenced using the primers Another doubtful record from the Canary Is. is presented by Hochkirch (2001). We used the P. canariensis (Saussure, 1884). Reexaminations HotMasterMix (Eppendorf, Hamburg, Germany) of museum collections (Natural History Museum, and the HotStarTaq Master Mix kit (Qiagen) for London and Zoölogisch Museum, Amsterdam) amplification. The PCR product was purified using revealed that this species has often been confused a Qiaex II gel extraction kit (Qiagen) according , with S. savignyi (Saussure, 1884), S. sublaevis to the manufacturer s protocol. Sequencing was (Bolívar, 1908), and S. azurescens (Rambur, performed with the Big Dye sequencing kit (Perkin 1838). Uvarov (1930) noted that the epitheton Elmer, Cheshire, UK) for sequencing reactions of P. canariensis is misleading as the types of run on a Perkin-Elmer ABI automated sequencer. this species were collected from Cape Verde and DNA sequences were corrected and aligned by not from the Canary Is. Johnsen (1974) used eye. We did not include ambiguous data from differences in the stridulatory mechanisms to the beginning and end of the fragment in the distinguish between S. canariensis and S. savignyi, analysis, resulting in a 1047-bp-long dataset. but our own studies show that these traits seem Sequences were deposited in GenBank under to be rather variable even within S. savignyi (MH the accession numbers EU266710-46. We used and AH unpubl. data). We found specimens with 3 different methods to infer a gene tree from serrate intercalary veins, with stridulatory cross our data. First, we used Bayesian Inference as veinlets between the radius and media (“knots”), implemented in MrBayes 3.1.1 (Ronquist and and with both traits combined, which will be Huelsenbeck 2003). The best-fitting substitution addressed in a future paper. model was chosen through the Akaike information During a field trip in 2006 (Husemann and criterion (AIC) as implemented in MrModeltest Hochkirch 2008), we obtained material of a new 2.2 (Nylander 2004). We ran the Monte Carlo species of Sphingonotus, which we originally Markov chain for 106 generations, sampling every identified as W. pachecoi (Bolívar, 1908). We 1000 generations. We discarded 1000 trees incorporated some specimens in a phylogenetic as burn-in, after checking that the chains were analysis to reconstruct the colonization pattern of stationary and convergent. Support of the nodes the Canary Is. (Fig. 1). Surprisingly, specimens was assessed by the posterior probabilities of from Fuerteventura represented a genetically reconstructed clades as estimated by MrBayes distinct lineage, while specimens from Lanzarote 3.1 (Ronquist and Huelsenbeck 2003). Using the were genetically rather similar to S. azurescens Minimum Evolution method, we visualized the from northern Africa and S. sublaevis, an endemic phylogenetic relationships among the samples species from Gran Canaria. As the type locality based on their pairwise Kimura 2-parameter of W. pachecoi is Lanzarote, the individuals distances as implemented in MEGA 3.1 (Kumar et from Fuerteventura had to belong to a hitherto al. 2004). Maximum-parsimony (MP) analysis was undescribed species, which is described below. performed in PAUP 4.0b10* (Swofford 2002), using a heuristic search (with TBR branch swapping). The confidence of the nodes was evaluated by MATERIALS AND METHODS bootstrapping the matrix 1000 times (Felsenstein 1985). Phylogenetic analysis Morphological analyses In total, 37 specimens of all Canarian Sphingonotus species as well as some We examined 5 males and 3 females northwestern African and European relatives were collected from Fuerteventura (see below) and obtained from 2002 to 2007 and stored either in a compared them with other Canarian as well as freezer or in absolute ethanol (Table 1). We chose with some North African and European species. the Oedipodinae species Oedipoda caerulescens We dissected the genitalia of a male of each (Linnaeus, 1758) and Sphingonotus scabriculus species according to the method described by Stål, 1876 from Namibia as outgroups. DNA was Hochkirch (2001). Moreover, we dissected the extracted from thoracic or femoral muscle tissue forewings of some specimen and examined the using the DNEasy tissue kit (Qiagen, Hilden, stridulatory apparatus with a digital scanning , Germany) following the manufacturer s protocols. electron microscope (SEM; Zeiss DSM 962, Hochkirch and Husemann - Review of Canarian Sphingonotini 497 Oberkochen, Germany). Measurements were Netherlands (ZMA); Museo de Ciencias Naturales taken with an ocular micrometer in the microscope. de Tenerife, Santa Cruz de Tenerife, Spain (MCN); Abbreviations for depositories are as follows: Departamento Biología Animal, Univ. de La Museum Alexander Koenig, Bonn, Germany Laguna, Tenerife, Spain (ULT); Academy of Natural (ZFMK); Zoölogisch Museum Amsterdam, The Sciences, Philadelphia, PA, USA (ANSP); and K233 Sph ru Ma K23 Sph ru GC K99 Sph ru LG 85/0.99 K255 Sph ru E S. rubescens K2 Sph ru Lanz 57/0.59 K71 Sph ru LP K88 Sph ru TF 99 K241 Sph wi TF S. willemsei 1.0 K240 Sph wi TF 100 96 K263 Sph co Co S. corsicus 0.99 1.0 K262 Sph co Co 66 99 K176 Sph ca Nu S. caerulans 0.95 1.0 K174 Sph ca Bo K34 Wer pi TF 50/0.67 100 1.0 K84 Wer pi TF W. picteti K38 Wer pi TF 93 K14 Wer gu GC W. guancha 0.53 K5 Wer ru Lanz W. rugosa 98 K228 Pse fi Ma 100 P. finotianus 1.0 K74 Pse fi Tu 0.86 K354 Sph nov Fue 98 100 K353 Sph nov Fue S. nov. sp. 0.86 0.99 K352 Sph nov Fue 60/ 0.51 K349 Sph sa Fue K87 Sph sa TF S. savignyi K163 Sph sa Go 76/ K4 Wer pa Lanz 0.99 K3 Wer pa Lanz W. pachecoi K76 Wer pa Lanz 88 K254 Pse az Tu 0.98 P. azurescens K222 Pse az Ma K219 Pse az Ma P. azurescens K20 Sph su GC K63 Sph su GC S. sublaevis K11 Sph su GC K215 Sph sc Na S. scabriculus K49 Oe cae D O. caerulescens 0.02 Fig. 1. Minimum Evolution tree based on pairwise Kimura 2-parameter distances of the NADH-dehydrogenase subunit 5. Values above the branches represent bootstrap values based on 1000 bootstrap replicates (only values > 50 are shown). Values behind the slash or below the branches represent percentage posterior probabilities from the Bayesian analysis.