Veromessor chicoensis

Veromessor chicoensis
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Myrmicinae
Tribe:	Stenammini
Genus:	Veromessor
Species:	V. chicoensis
Binomial name
Veromessor chicoensis Smith, M.R., 1956 Specimen Labels

Little is known about the biology of V. chicocensis. Workers are diurnal foragers during spring and summer months, but shift to crepuscular foraging at temperatures of about 30° C and forage nocturnally at temperatures of 35° C (Hamm & Kamansky, 2009). Previous indications were that V. chicoensis workers were solitary foragers based on “no conspicuous foraging trail was observed” (M. Bennett, pers. comm.). However, the senior author observed two colonies in Tulare County in early October 2018 that were foraging in a narrow 5–10 m long column during mid-morning. Colonies appeared to contain about 1,000–2,000 workers (R.A. Johnson, pers. obs). Workers are polymorphic. Nests are placed in hard clay soils. Veromessor chicoensis is sometimes sympatric with Veromessor andrei, whereas it appears to occur without V. andrei at other locations, especially those with marginal, thin soil habitats (P.S. Ward, pers. comm.). (Johnson et al., 2022)

Identification

Worker

This species is uniquely characterized by the following combination of features (Johnson et al., 2022):

1. Head and mesosoma orangish-red or orangish-brown to reddish-brown, gaster blackish-orange to blackish
2. Medial lobe of clypeus lacking medial groove but with several lateral longitudinal rugae, medial lobe not thick and protuberant in profile, not elevated above lateral lobes in frontal view
3. Mandibles with 8 teeth
4. Scape narrowest near midlength,
5. MOD distinctly less than OMD, OI < 23.0
6. Cephalic dorsum weakly shining between fine longitudinal rugae, usually becoming somewhat reticulate on posterior margin; antennal fossa sometimes with several concentric rugae
7. Psammophore poorly developed; ventral surface of head capsule with scattered straight or evenly curved hairs, but J-shaped hairs mostly absent
8. Pronotum with fine, wavy to irregular, longitudinal rugae; sides of pronotum with longitudinal rugae that sometimes fade to disappear posterad, becoming granulate posterad; mesonotum with wavy to weakly irregular longitudinal rugae; mesopleura with wavy to irregular rugae that traverse longitudinally to posterodorsally, rugae sometimes with lateral branches, interrugae moderately to strongly granulate
9. Propodeal spines short, acuminate, length less than distance between their bases; infraspinal facet and propodeal declivity strongly coriarious or rugose, weakly dull to weakly shining
10. Metasternal process large, higher than long with broadly rounded apex; anterior and posterior surfaces decline steeply; weakly translucent in profile (Figures 6A, 15–16)

• 

  Johnson et al, 2022, Fig. 6. Photographs of the four categories of increasing psammophore development for species of Veromessor based on number and distribution of long J-shaped hairs on the ventral surface of the head capsule (= hypostomal region) (see text). Photograph of: (A) V. chicoensis (CASENT0923125)—J-shaped hairs mostly absent with scattered straight or evenly curved hairs, (B) V. andrei (CASENT0923140)—J-shaped hairs present but not arranged in a distinct row, usually mixed with straight or evenly curved hairs, (C) V. chamberlini (CASENT0761101) and (D) V. smithi (CASENT0923131)—J-shaped hairs present, arranged in a V-shaped row which does not reach the posterior part of the lateroventral margin of head capsule, usually mixed with straight or evenly curved hairs, and (E) V. pergandei (CASENT0923124)—J-shaped hairs present, many long J-shaped hairs arranged in a distinct row around the outer margin of the ventral region of the head capsule. Photographs by Wade Lee from www.AntWeb.org.
• 

  Johnson et al., 2022, Fig. 15. Photograph of Veromessor chicoensis minor worker: (A) frontal view of head, (B) lateral view of body, and (C) dorsal view of body (LACMENT359751). Photographs by Michele Esposito from www.AntWeb.org.
• 

  Johnson et al., 2022, Fig. 16. Photograph of Veromessor chicoensis major worker: (A) frontal view of head, (B) lateral view of body, and (C) dorsal view of body (LACMENT359767). Photographs by Michele Esposito from www.AntWeb.org.

Queen

This caste is diagnosed by the following combination of features (Johnson et al., 2022):

1. Head and mesosoma dark orangish to orangish-brown with parts of mesopleura and mesoscutum blackish-orange to blackish; gaster blackish orange
2. Medial lobe of clypeus with strong submedial and sublateral rugae
3. Mandibles with 8 teeth
4. Dorsal base of scape weakly flattened; maximum basal width of scape greater than maximum preapical width
5. MOD less than to greater than OMD
6. Cephalic dorsum weakly shining between fine, wavy to irregular, longitudinal rugae; medial rugae diverging toward posterior corners, rugae becoming strongly irregular to weakly rugoreticulate along posterior margin
7. Psammophore poorly developed
8. Sides of pronotum weakly shining between fine, longitudinal rugae; mesoscutum with fine, longitudinal rugae; mesoscutellum smooth and shining; anepisternum weakly shining between fine longitudinal rugae; katepisternum moderately shining between slightly more widely spaced rugae
9. Sides of propodeum weakly shining and densely punctate-granulate between longitudinal and oblique rugae; propodeal spines triangular, acuminate, length less than distance between their bases; infraspinal facet and propodeal declivity with fine transverse rugae, interrugae weakly dull to weakly shining
10. Metasternal process large, higher than long with broadly rounded apex; anterior and posterior surfaces decline steeply; weakly translucent in profile (Figure 17)

• 

  Johnson et al., 2022, Fig. 17. Photograph of Veromessor chicoensis alate queen: (A) frontal view of head, (B) lateral view of body, and (C) dorsal view of body (CASENT0922823). Photographs by Michele Esposito from www.AntWeb.org.

Male

This caste is diagnosed by the following combination of features (Johnson et al., 2022):

1. Head and mesosoma dark brown to dark blackish-brown; gaster a lighter medium brown
2. Clypeus weakly convex in profile; anteromedial margin weakly concave, with a weak medial longitudinal carina
3. Mandibles with 1–3 teeth basad of preapical tooth
4. Anterior ocellus well above level of top of eyes
5. Mesopleura weakly shining, moderately to strongly lineopunctulate-lineogranulate, sometimes with rugae along posterior margin
6. Propodeum contiguously punctategranulate and with scattered, irregular, fine rugae; propodeal spines cariniform to knoblike and inconspicuous,
7. Metasternal process prominent and acute to elongate and narrowly rounded
8. Subpetiolar process short, elongate-triangular to spiniform, apex acuminate to rounded (Figures 1C, 18).

• 

  Johnson et al., 2022, Fig. 1. Photographs of the metasternal process on males of Veromessor: (A) V. andrei (CASENT4010823), (B) V. chamberlini (UCRC_ENT00500152), (C) V. chicoensis (CASENT0869853), (D) V. julianus (LACMENT359792), (E) V. lariversi (CASENT0761204), (F) V. lobognathus (LACMENT363986), (G) V. pergandei (CASENT0869850), (H) V. pseudolariversi (CASENT0869851), (I) V. smithi (LACMENT364071), and (J) V. stoddardi (LACMENT364102). Photographs by Robert Johnson from www.AntWeb.org.
• 

  Johnson et al., 2022, Fig. 18. Photograph of Veromessor chicoensis male: (A) frontal view of head, (B) lateral view of body, and (C) dorsal view of body (LACMENT359769). Photographs by Michele Esposito from www.AntWeb.org.

Identification Notes

The poorly developed psammophore and short propodeal spines distinguish workers of V. chicoensis from all congeners except the closely related Veromessor stoddardi. Veromessor chicoensis is distinguished by: (1) antennal scape narrowest near mid-length; maximum basal width of scape greater than maximum preapical width, and (2) in dorsal view, rugae on mesonotum similar to those on pronotum, interrugae weakly dull. In V. stoddardi: (1) antennal scape narrowest immediately distad of basal bend; maximum basal width of scape about equal to maximum preapical width, and (2) in dorsal view, the mesonotum usually smooth and shining, rugae distinctly weaker than those on pronotum. (Johnson et al., 2022)

• 

  Johnson et al., 2022, Fig. 19. Bivariate plots for workers, queens, and males of Veromessor chicoensis and V. stoddardi: (A) head width versus head length and (B) head width versus mesosoma length. Sample size for each caste is in parentheses.
• 

  Johnson et al., 2022, Fig. 55. Bivariate plots for workers showing head width versus head length (left panels) and mesosoma length versus pronotum width (right panels) for ecologically similar pairs of small colony species of Veromessor: V. lariversi and V. pseudolariversi (A–B), V. lobognathus and V. smithi (C–D), V. chicoensis and V. stoddardi (E–F). Note that the x-axis scale is the same for all panels on the left and for all panels on the right so that sizes can be compared directly across species groups. Sample size for each species is given in parentheses.
• 

  Johnson et al., 2022, Fig. 53. Eye area (mm2) (A), facet number (B), and mean facet diameter (μm) (C) for pale and dark colored species of Veromessor. Two species are pale (V. lariversi, V. pseudolariversi—open symbols and regular font), while the other eight species are dark (filled symbols and bold font). For each species, number of workers examined and number of colonies they were derived from is given in parentheses. Significant differences (P < 0.05) among species are denoted after each species name by the letters a–g: a > b > c > d > e > f > g; the three sets of letters for each species correspond to panels A, B, and C, respectively. Groupings are based on univariate F tests within MANCOVA followed by pairwise comparisons using a least significant differences test (see also Johnson & Rutowki, 2022).
• 

  Johnson et al., 2022, Fig. 56. Queen body size across species of Veromessor based on average head width, average head length, and average mesosoma length. Species are ordered (left to right) from shortest to longest mesosoma length.

Keys including this Species

• Key to Veromessor species

Distribution

This species occurs at elevations from 60–660 m in the California interior chaparral and woodlands and California Central Valley grasslands ecoregions, as defined by Olson et al. (2001) (Johnson et al., 2022).

Veromessor chicoensis appears to be restricted to the Sacramento and San Joaquin Valleys in northern and central California and in low mountains along the northwestern edge of the Sacramento Valley. Veromessor chicoensis probably occurred throughout most to all of the San Joaquin Valley historically given a 1947 record of one male from Tulare County, and more recent collections from Tulare County near vernal pools in Pixley National Wildlife Refuge and James K. Herbert Wetland Prairie Preserve (Hamm & Kamansky, 2009). The senior author collected two colonies in the James K. Herbert Wetland Prairie Preserve in October 2018, but this species was not located during a brief survey at Pixley National Wildlife Refuge at that time. Moreover, the current range of V. chicoensis in the San Joaquin Valley appears to be limited to isolated properties and refuges that have not been developed for agriculture (Hamm & Kamansky, 2009). (Johnson et al., 2022)

• 

  Johnson et al., 2022, Fig. 10. Geographic distribution of: (A) Veromessor andrei, (B) V. chamberlini, and (C) V. chicoensis. The larger black circle in each panel denotes the type locality. The northernmost locale for V. chamberlini was given only as Santa Clara County, and we have placed this locale near the center of the county.

Latitudinal Distribution Pattern

Latitudinal Range: 39.728494° to 38.35°.

North Temperate	North Subtropical	Tropical	South Subtropical	South Temperate

• Source: AntMaps

Distribution based on Regional Taxon Lists

Nearctic Region: United States (type locality).

Distribution based on AntMaps

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.

Biology

The following notes are provided by Johnson et al. (2022).

Mating flights have not been observed, but sexuals have been collected from May 14 through August 28, suggesting that mating flights start early in the summer and last for an extended period. Nothing is known about colony founding, but head width and head length of queens of V. chicoensis are about the same as medium-sized workers, whereas mesosoma length is greater in queens than in workers (Figure 19). Their small size infers that founding queens of V. chicoensis are semiclaustral, probably obligate foragers.

Veromessor chicoensis is also unusual because dealate queens sometimes occur in foraging columns (one foraging dealate queen collected by the senior author on October 2, 2018). Foraging by dealate queens is common in V. andrei, but this is the first such record in V. chicoensis (see above discussion under Veromessor andrei).

Gland chemistry has not been examined in V. chicoensis. Like other small-colony congeners, workers of V. chicoensis have a small pygidial gland reservoir and lack a textured tergal cuticle (Hölldobler et al., 2013).

The Pacific Coast horned lizard (Phrynosoma cornutum) preys upon V. chicoensis, and it sometimes is the main component in the diet of these lizards (Hamm & Kamansky, 2009).

Castes

Images from AntWeb

Worker. Specimen code casent0217879. Photographer Estella Ortega, uploaded by California Academy of Sciences.	Owned by CAS, San Francisco, CA, USA.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

• chicoensis. Veromessor stoddardi subsp. chicoensis Smith, M.R. 1956a: 37 (w.) U.S.A. (California).
  • Type-material: holotype worker, 20 paratype workers.
  • Type-locality: U.S.A.: California, Butte County, Chico, 2.iii.1954, 450 ft, Sec.17, T22N, R2E (A. Wenner).
  • Type-depository: USNM.
  • Combination in Messor: Bolton, 1995b: 253;
  • combination in Veromessor: Ward, et al. 2015: 73.
  • Subspecies of stoddardi: Smith, M.R. 1958c: 120; Smith, D.R. 1979: 1365; Bolton, 1995b: 253.
  • Status as species: Johnson, R.A. 2000: 96 (in list); Ward, 2005: 66.
  • Distribution: U.S.A.

Type Material

• Holotype worker [USNM: USNMENT00529087], USNM#62959, 8 paratype workers [USNM], 6 paratype workers [LACM], UNITED STATES, California: Butte County, Chico (T22N, R2E, Section 17), 450’, 2 March 1954 (Adrian Wenner) (examined by Johnson et al., 2022).

Johnson et al. (2022) - Smith (1956) described V. stoddardi chicoensis from a holotype and 20 paratype workers and indicated specimens were deposited at USNM. We examined nine type workers (on three pins) from USNM. All nine had type labels USNM #62959, but none of these specimens had a holotype label. Two pins had a USNM Paratype No. 62959 label, and one pin had a USNM Type No. 62959 label; the latter pin also had a black dot on the uppermost triangle, and this specimen is presumed to be the holotype. Consequently, we placed a holotype label on this worker.

References

• Johnson, R.A., Borowiec, M.L., Snelling, R.R., Cole, A.C. 2022. A taxonomic revision and a review of the biology of the North American seed-harvester ant genus Veromessor (Hymenoptera: Formicidae: Myrmicinae). Zootaxa 52061, 1-115 (doi:10.11646/zootaxa.5206.1.1).
• Plowes, N.J.R., Johnson, R.A., Holldobler, B. 2013. Foraging behavior in the ant genus Messor (Hymenoptera: Formicidae: Myrmicinae). Myrmecological News 18, 33-49.
• Ward, P.S., Brady, S.G., Fisher, B.L. & Schultz, T.R. 2014. The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology, DOI: 10.1111/syen.12090

References based on Global Ant Biodiversity Informatics

• Johnson R. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
• Smith M. R. 1956. A key to the workers of Veromessor Forel of the United States and the description of a new subspecies (Hymenoptera, Formicidae). Pan-Pacific Entomologist 32: 36-38.