• Tweet this page
• Like this page
• Email this page
• Share this page

• Home»
• Publications »
• Books & monographs »
• Fauna of New Zealand Series »
• Extracts »
• FNZ44: Lycosidae »
• Phylogenetic analysis

FNZ 44 - Lycosidae (Arachnida: Araneae) - Phylogenetic analysis

Vink, CJ 2002. Lycosidae (Arachnida: Araneae). Fauna of New Zealand 44, 94 pages.
( ISSN 0111-5383 (print), ; no. 44. ISBN 0-478-09347-0 (print), ). Published 23 Dec 2002
ZooBank: http://zoobank.org/References/3410DFC9-CB06-42DB-8B88-AD93A35AE4D2

Phylogenetic analysis

Methods. A reconstruction of the phylogenetic history of New Zealand lycosid species was attempted using a cladistic analysis of morphological characters. The analysis included all species of Anoteropsis and the three New Zealand Artoria species as the outgroup taxa. Allotrochosina schauinslandi, Geolycosa tongatabuensis, and Venatrix goyderi were not included in the analysis as they were polyphyletic to Anoteropsis and Artoria (see Vink et al. 2002). Artoria appears to be the sister genus to Anoteropsis (Vink et al. 2002) and was, therefore, selected as an outgroup (see Watrous & Wheeler 1981 and Maddison et al. 1984 for a discussion of outgroups). Notocosa bellicosa was considered too distant from Anoteropsis and Artoria (see Vink et al. 2002) to be a meaningful outgroup taxon in the analysis. Although it has been argued that the outgroup need not be the sister group of the ingroup (Nixon & Carpenter 1993), the inclusion of N. bellicosa in the data matrix lowered the resolution of the cladogram.

Eight morphological characters used in the analysis were from male pedipalpal morphology. Male genitalia are commonly used in spider phylogenetic analyses (e.g., Coddington 1990, Hormiga 1993, Griswold 2001). Characters were also taken from somatic morphology (seven), female genitalia (six), and ecology (one). Distinct gaps in ratios and measurements were used to separate character states.

Character list

1. Anterior eye row: (0) slightly procurved or straight (Fig. 3, 5); (1) strongly procurved (Fig. 4).
2. Anterior eye row: (0) no more than one anterior lateral eye width apart; (1) more than one anterior lateral eye width apart.
3. Retromarginal cheliceral teeth: (0) three - none reduced; (1) three - proximal reduced.
4. Scopulae on tarsi and metatarsi I and II: (0) absent; (1) weak (comparatively large spaces between adjacent scopulae); (2) intermediate (small spaces between adjacent scopulae); (3) dense (an almost solid mass, scopulae contiguous).
5. White pubescence below PME: (0) absent; (1) present.
6. Cymbium/tibia length ratio: (0) more than 2.1; (1) 2.0-1.5; (2) less than 1.4.
7. Minimum adult body length: (0) less than 8 mm; (1) greater than 9 mm.
8. Basoembolic apophysis: (0) without spur; (1) with short spur (small, sclerotised bump); (2) with long spur (at least ¼ length of median apophysis).
9. Lobe at base of terminal apophysis: (0) no lobe; (1) weak lobe; (2) rounded lobe; (3) tooth-like lobe.
10. Length of median apophysis after bend: (0) without bend; (1) median apophysis with weak bend; (2) less than or equal to length before bend; (3) longer than before bend.
11. Median apophysis dorsoventrally flattened: (0) no; (1) partially.
12. Median apophysis with basal spur: (0) absent; (1) present.
13. Median apophysis shape after bend: (0) absent; (1) pointing anteriorly (e.g., Fig. 11); (2) wave-like (e.g., Fig. 25); (3) straight (e.g., Fig. 18); (4) pointing posteriorly (e.g., Fig. 24).
14. Median apophysis tip: (0) rounded; (1) mesially directed tooth; (2) one laterally directed tooth; (3) two laterally directed teeth; (4) one laterally directed tooth and blunt protrusion below tooth.
15. Tip of terminal apophysis: (0) pointed; (1) rounded; (2) multifaceted.
16. Epigynal hoods: (0) absent; (1) shallow; (2) regular; (3) deep.
17. Posterior lip: (0) absent; (1) present.
18. Median septum: (0) not raised; (1) raised and unsclerotised; (2) raised and sclerotised.
19. Bends in internal genitalia: (1) 1; (2) 2; (3) 3; (4) 4; (5) 5 or more.
20. Epigyne raised either side of septum: (0) no; (1) yes.
21. Genitalia lateral sclerites: (0) absent; (1) present.
22. Habitat: (0) grassland - including open scrub and swamp; (1) stony or sandy - including riverbed, scree and beach; (2) forest - including forest litter.

Results. The data matrix (Table 1) was analysed with PAUP* version 4.0b8 (Swofford 2001) using the branch and bound search, which finds all most parsimonious trees. Characters 8, 16, and 19 were ordered. Polarity was not assigned to any characters and all characters had equal weighting a priori, as there was no obvious weighting scheme. All 22 characters were phylogenetically informative. Characters that were autapomorphic or appeared to be homoplasious (e.g., colour pattern, which varied intraspecifically) were excluded a priori from the analysis. Bootstrap values (Felsenstein 1985) were calculated from 1000 replicate parsimony analyses using the closest addition sequence of the taxa and the heuristic search option in PAUP*.

Table 1. Character state distribution matrix for phylogenetic analysis of Anoteropsis.

Parsimony analysis produced one most parsimonious tree (Text-fig. 2), with a length of 77 steps, a consistency index of 0.571 and a retention index of 0.761.

Relationships. There is strong (>75%) bootstrap support for A. hilaris and A. ralphi as sister species. Anoteropsis alpina, A. blesti, A. hallae, A. montana, and A. westlandica appear to be basal within Anoteropsis and all other species form a derived clade, with strong bootstrap support. Anoteropsis arenivaga, A. canescens, and A. urquharti are morphologically similar and form an unresolved trichotomy.   

The Lycosidae is a morphologically conservative family, which makes it difficult to generate large morphological datasets for phylogenetic analysis. Because of this the data matrix was relatively small (fewer characters than taxa); however, there was still good resolution and bootstrap support within the tree. Further resolution through additional phylogenetic analyses was carried out on Anoteropsis species using data from partial sequences from cytochrome c oxidase subunit I and NADH dehydrogenase subunit I plus tRNALeu(CUN) DNA (Vink & Paterson, in prep.). The phylogenies inferred from these molecular data were significantly similar (p<0.001) to the phylogeny presented here.

Purchase this publication

• Purchasing details