Kingdom Animalia (Animals)
Phylum Arthropoda (Arthropods)
Subphylum Hexapoda (Hexapods)
Class Insecta (Insects)
Order Phasmida (Walkingsticks)
, of which 20 in our area(2)
There are also at least two more known but undescribed taxa...one using Limber Pine (Pinus flexilis
) as host plant; and one using Sargent Cypress (Hesperocyparis sargentii
) as host plant.
Body length: Males: 11-20 mm ; Females: 14-28 mm
The species distribution map below (see under "Range
"), and the host plant list below (see under "Food
" ), can help narrow ID possibilities (though neither are necessarily complete or entirely accurate).
But dependable species ID requires study of the shape of externally visible structures of the terminalia, especially of the male (for non-parthenogenetic spp.)...in conjunction with location, host plant, color and markings.
Note to photo contributors: ID will be facilitated by providing detailed images of terminalia (dorsal view for males; dorsal, ventral, and lateral for females), as well as full-body dorsal views indicating overall color and markings.
Males have asymmetric terminalia with two variously lobed cerci ("claspers"), and a more-or-less ribbon-like projection between them (positioned next to the right cercus and called the "intradextral process"). Figures appear here
. Keys refer to left and right cerci using the latin terms "sinistral" and "dextral", respectively.
Female terminalia are symmetric with somewhat longer, more uniformly cylindrical cerci.
A one-page visual summary of diagnostic diagrams of the terminalia for all 21 currently described species appears on pg. 61 of Vickery & Sandoval (2001).
An online key adapted from Vickery (1993) can be found here
...but it covers only 12 species, omitting the 9 species described after 1993.
A remarkable aspect of Timema is the existence of a number of parthenogenetic species consisting solely of asexually reproducing females.
Females in bisexual species are rarely seen alone, due to male "mate guarding", in which males ride on the backs of females (for periods up to several days). Males are noticeably smaller than females in body size. According to Sandoval & Vickery (1996), a field observation of 5 or more adult females with no males present is a likely indicator of an asexual species.
To each asexual species, there corresponds a closely related sexual species in which the females of the two are visually indistinguishable. These species pairings are as follows:
T. genevievae and T. podura;
T. tahoe and T. bartmani;
T. douglasi and T. poppensis;
T. shepardi and T. californicum;
T. monikensis and T. cristinae;
sw. US and Mexico...primarily in CA, but also in s. OR, far w. NV, far s. NV, AZ, and Baja California.
Note that the range map below is not exact and should be cross-referenced with the map of point records here
, as well as collection records from the literature
In particular, the map below under-represents distributions of some species. For instance:
1) T. californicum has records north of San Francisco Bay in Marin Co.;
2) T. poppenis and T. douglasii are recorded from south of San Francisco Bay in the Santa Cruz Mnts;
3) T. genevievae has records in the serpentine areas of eastern Lake and Napa counties; etc.
So use the map from Wikipedia below as a helpful but rough guide only!
Map from Wikipedia, based on map from pg. 1473 of Law & Crespi (2002)
On foliage, twigs, or branches of host shrubs or trees...or on the ground, where they drop to upon disturbance. Host plants mostly associated with chaparral; some with woodlands or forest (e.g. douglas fir, redwood).
Green morphs tend to rest on leaves; brown to gray morphs on stems, branches or ground.
Unstriped morphs are usually associated with broad-leaved host plants (e.g. oaks, ceanothus, manzanita, etc.). Striped morphs are usually associated with host plants having needle-like leaves (e.g. chamise, douglas fir, redwood, etc.).
Coloration, stripes, and other markings serve as camouflage, and are adaptations driven by selection pressure due to predation by visually-oriented birds and lizards. For interesting details, see Sandoval (1994), Sandoval & Crespi (2008), and this course handout & worksheet
Timema typically feed on leaves at night and rest by day on leaves or stems of their host plants. The following list gives known hosts for each species:
T. bartmani: white fir (Abies concolor)
T. boharti: manzanita (Arctostaphylos spp.), ceanothus (Ceanothus spp.), chamise (Adenostoma fasciculatum), oak (Quercus spp.)
T. californicum: manzanita (Arctostaphylos spp.), ceanothus (Ceanothus spp.), oak (Quercus spp.), mountain mahogony (Cercocarpus spp.), toyon (Heteromeles arbutifolia)
T. chumash: ceanothus (Ceanothus spp.), oak (Quercus spp.), mountain mahogony (Cercocarpus spp.)
T. coffmani: juniper (Juniperus spp.)
T. cristinae: ceanothus (Ceanothus spp.), chamise (Adenostoma fasciculatum), toyon (Heteromeles arbutifolia)
T. dorotheae: ceanothus (Ceanothus spp.)
T. douglasi: douglas fir (Pseudotsuga menziesii)
T. genevievae: chamise (Adenostoma fasciculatum)
T. knulli: coast redwood (Sequoia sempervirens), ceanothus (Ceanothus spp.)
T. landelsensis: manzanita (Arctostaphylos spp.)
T. monikensis: ceanothus (Ceanothus spp.), mountain mahogony (Cercocarpus spp.)
T. morongensis: wild buckwheat (Eriogonum spp.)
T. nakipa: manzanita (Arctostaphylos spp.), ceanothus (Ceanothus spp.), chamise (Adenostoma fasciculatum)
T. nevadense: pinyon pine (Pinus monophylla); juniper (Juniperus spp.)
T. petita: ceanothus (Ceanothus spp.)
T. podura: chamise (Adenostoma fasciculatum), ceanothus (Ceanothus spp.), oak (Quercus spp.), mountain mahogony (Cercocarpus spp.)
T. poppensis: douglas fir (Pseudotsuga menziesii), coast redwood (Sequoia sempervirens)
T. ritensis: juniper (Juniperus spp.)
T. shepardi: manzanita (Arctostaphylos spp.)
T. tahoe: white fir (Abies concolor)
In most species, eggs are dropped from branches of host plants.
In all species, during egg-laying, females defecate previously-ingested soil, and then use their cerci to firmly cover and pack the eggs with the soil. It's hypothesized this may prevent desiccation, reduce parasitism by micro-hymenoptera, and perhaps provide some protection from fire, which is common in chaparral habitats where Timema are found. Moreover, soil is found in the gut of all newly hatched larvae and Sandoval has observed that laboratory-hatched eggs of all species survived only when provided with soil from their native locality in the first few days after hatching. It is thought that the soil provides offspring access to microbial symbionts for digestion of cellulose. Sandoval states that soil ingestion may also reduce toxicity effects from host plant secondary compounds on the larvae.
Parasitism by tachnid flies has been observed for the larvae of T. douglasi, T. cristinae, and T. californicum.
is a genus of small, stout, wingless walking sticks. It is so distinctive that it is the only genus in the entire suborder Timematodea
, and it is an ancient group which is phylogenetically basal to the rest of the walking stick order Phasmida. (See Tree of Life website
and illustrated cladogram here
A fascinating aspect of Timema is the existence of five parthenogenetic species consisting of asexually reproducing females. These are: T. douglasi, T. genevievae, T. monikensis, T. shepardi, and T. tahoe. Populations of these species consist (almost) entirely of females (a few anomalous males have been collected in T. monikenesis and T. shepardi). Eggs laid by field collected females of these species hatch in the lab to females, which in turn lay eggs without mating that again yield females. In recent years researchers have concluded that some of these species (e.g. T. genevievae) have been asexually reproducing for over a million years. In general, most of the parthenogenetic species have a more northerly distribution than their corresponding sexual species. Various hypotheses have been explored to explain this , e.g. positing dispersal advantages for parthenogens.
T. genevievae, described by Rentz in 1978, was the first Timema species recognized as parthenogenetic. Rentz noted that while other species known at the time emitted an acrid odor, T. genevievae did not. David Weissman suggested that the acrid odor may be associated with mating-related pheromones, which might be unnecessary...and thus lost...in a long-time asexual species. (Vickery 1993, pg. 678).
The discovery and description of Timema species is interesting. The first species (T. californicum) was described from the Santa Cruz Mountain of California in 1895, and the second (T. chumash) was described from Los Angeles in 1920 by Hebard, who opined that they constituted the most "aberrant" group within the walking sticks. By 1966 eight species had been described from California and adjacent areas of southern Nevada and central Arizona. In 1978 the first parthenogenetic (= all female, asexually reproducing) species was described from southeast of the San Francisco Bay (T. genevievae). Between 1993 and 2001, twelve more species were described, four of which are parthenogenetic. More known species currently await description, and others likely await discovery.
D. Arbuthnott, M. G. Elliot, M. A. McPeek, & B. J. Crespi (2010). Divergent patterns of diversification in courtship and genitalic characters of Timema
walking-sticks. J. Evolution Biol. 23(7) 1399:1411 (PDF
Bartman, G. & Brock, P. D. (1995). Observations of the appearance and behavior of species of the stick-insect genus Timema
Scudder (Phasmida: Timematodea). Bulletin of the Amateur Entomologist's Society, 54:197-203 (Full Text
Hebard, M. (1920) The genus Timema
Scudder, with the description of a new species, (Orthoptera, Phasmidae, Timeminae). Entomological News, 31 126-132 (Read at BHL
Hebard, M. (1937) Studies in Orthoptera [...]:Notes and a new species of Timema
[...]. T. Amer. Entomol. Soc, 63 (3), pp. 347-350. (Read online
Law, J. H. & B. J. Crespi (2002). The evolution of geographic parthenogenesis in Timema
walking-sticks. Molecular Ecology 11: 1471–1489 (PDF
Maderspacher, Florian (2011). Asexuality: The Insects that Stick With It. Current Biology, 21 (13) pp. 495-497 (Read online here
Rentz, D. C. F. (1978). A new parthenogenetic Timema from California. Pan-Pacific Entomologist, 54 (3), 173:177.
Sandoval, C.P. (1994). Differential visual predation on morphs of Timema cristinae (Phasmatodea, Timemidae) and its consequences for host range. Biol. J. Linn. Soc., 52, pp 341-356.
Sandoval, C. P. and Nosil, P. (2005). Counteracting selective regimes and host preference evolution in ecotypes of two species of walking-sticks. Evolution 59:2405-2413 (Full Text
Sandoval, C.P. and Vickery, V.R. (1996). Timema douglasi (Phasmatoptera:Timematodea), a new parthenogenetic species from southwestern Oregon and northern California, with notes on other species. Can. Entomol. 128:79-84.
Sandoval, C. & B. J. Crespi (2008). Adaptive evolution of cryptic coloration: the shape of host plants and dorsal stripes in Timema
walking-sticks. Biol. J. Linn. Soc., 94, pp 1-5 (PDF
Schwander T. , H. Lee, B.J. Crespi (2011). Molecular evidence for ancient asexuality in Timema
stick insects. Curr. Biol., 21, pp. 1129–1134 (Full Text
Schwander T., Crespi B.J. (2009). Multiple direct transitions from sexual reproduction to apomictic parthenogenesis in Timema
stick insects. Evolution 63, 84-103. (PDF
Strohecker, H. F. (1966). New Timema from Nevada and Arizona. Pan-Pacific Entomologist, 42 (1) pp. 25-26.
Tinkham, E. R. (1942). A new californian species of Timema
(Phasmodea: Timemidae) with zoogeographical notes. Bull. S. Calif. Acad. Sci. 41(2):72-79. pl. 14 (Read at BHL
Vickery, V. R. (1993). Revision of Timema Scudder (Phasmatoptera: Timematodea) including three new species. Can. Entomol. 125:657–692.
Vickery, V. R. & Sandoval, C. P. (1997). Timema bartmani (Phasmatoptera: Timematodea: Timematidae), a new species from southern California. The Canadian Entomologist 129:933-936
Vickery, V. R. & Sandoval, C. P. (1998). Timema monikensis, Species Nov. (Phasmatoptera: Timemmatidea: Timematidae), a new parthenogenetic species in California. Lyman Entomological Museum and Research Laboratory, Note Number 22.
Vickery V. R. & C. P. Sandoval (1999a). Two new species of Timema
, One Parthenogenetic in California. J. Orthoptera Res. 8 (1) 45:47 (JSTOR
Vickery V. R. & C. P. Sandoval (1999b). Timema coffmani
(Phasmatoptera: Timematodea) a New Species from Arizona and Description of the Female of Timema ritensis
. J. Orthoptera Res. 8 (1) 49:52 (JSTOR
Vickery V. R. & C. P. Sandoval (2001). Description of three new species of Timema
and Notes on Three Other Species. J. Orthoptera Res. 10 (1) 53:61 (JSTOR
Article from Smithsonian
on Cristina Sandoval's research with T. cristinae
, and the light it sheds on the process of speciation.
Cristina Sandoval's Timema web page
: Images of 13 species, research project info, and bibliography...with some articles available as PDF's.
Tanja Schwander's Timema web page
: Color drawings of 8 species, interactive map with numerous records of most species, research project info, and bibliography...with some articles available as PDF's.
from Phasmida species file online, by P.D. Brock.