Use negative binomial regression with a logarithmic mean link, for example
library(MASS)
species_nb <- glm.nb(nspecies ~ altitude, link = log)
In the mean-size parametrization,
The extra parameter allows overdispersion that grows quadratically with the mean, unlike the proportional variance of Quasi-Poisson regression.
The printed assertion needs qualification: a negative-binomial model with fixed size is a generalized linear model. For known , its probability mass function is
With natural parameter , it has exponential family form
with dispersion one. A logarithmic link is permitted even though it is not the canonical link function. This is the fixed-size negative binomial generalized linear model.
What prevents a single ordinary fixed-family GLM fit in glm.nb is joint estimation of the unknown size : it changes the cumulant, carrier, and variance function, rather than merely multiplying a fixed variance function by a scalar dispersion. The routine alternates a GLM fit at fixed with size-parameter likelihood updates. Thus the intended distinction is between that enlarged fitting problem and a single fixed-family GLM, not a claim that negative-binomial regression can never be a GLM. The package documentation explicitly supports both conventions: stat.ethz.ch/R-manual/R-patched/library/MASS/html/glm.nb.html and stat.ethz.ch/R-manual/R-patched/library/MASS/html/negative.binomial.html.

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